OCCASIONAL PAPERS OF THE ■ No. 141, 104 pages, 142 figures. CALIFORNIA ACADEMY OF SCIENCES OCT 2 ism September 19, 1984 Systematics of Eusattus and Conisattus (Coleoptera; Tenebrionidae; Coniontini; Eusatti) By John T. Doyen University of California, Berkeley, California 94720 ^sm^y %DED\^ £> SAN FRANCISCO PUBLISHED BY THE ACADEMY ' OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES No. 141, 104 pages, 142 figures. September 19, 1984 Systematics of Eusattus and Conisattus (Coleoptera; Tenebrionidae; Coniontini; Eusatti) By John T. Doyen University of California, Berkeley, California 94720 ^DBI^ %DED\^ SAN FRANCISCO PUBLISHED BY THE ACADEMY TABLE OF CONTENTS Abstract iv Introduction 1 Acknowledgments 1 Materials and Methods 2 Biology 4 Higher Classification of Coniontini 6 Cladistic Relations Among Eusattus Species Groups 1 3 Relationships Within Species Groups 1 6 Distributional Patterns of Eusattus 1 9 Key to the Genera of Coniontini 25 Conisattus Casey 25 Conisattus rectus Casey 25 Eusattus LeConte 28 Key to the Species of Eusattus 29 Convexus Species Group 35 Eusattus convexus LeConte 35 minimus Doyen 38 nitidipennis LeConte 40 obliteratus Champion 41 Reticulatus Species Group 42 Eusattus araneosus (Blaisdell) 43 aridus Doyen 43 catalinensis Doyen 45 catavinus Doyen 46 cedrosensis Doyen 47 ceralboensis Doyen 48 cienegus Doyen 49 costatus Horn 50 erosus Horn 52 franciscanus Doyen 54 laevis LeConte 55 mexicanus Champion 56 planulus Doyen 57 pons Triplehorn 59 reticulatus (Say) 60 venosus Champion 62 Muricatus Species Group 63 Eusattus dilatatus LeConte 64 hirsutus Doyen 66 muricatus LeConte 67 phreatophilus Doyen 71 puberulus LeConte .! 73 Ciliatus Species Group 74 Eusattus arenarius Doyen 75 ciliatoides Doyen 75 ciliatus Horn 77 dubius LeConte 78 pallidus Doyen 83 vizcainensis Doyen 85 Depressus Species Group 86 Eusattus depressus Champion 87 secutus Horn 87 Rudei Species Group 89 Eusattus crypt icus Doyen 90 rudei Doyen 91 Robustus Species Group 92 Eusattus politus Horn 93 robustus LeConte 93 Difficilis Species Group 95 Eusattus difficilis LeConte 96 productus LeConte 97 Literature Cited 99 Appendices 100 Abstract Doyen, John T. Systematics of Eusattus and Conisattus (Coleoptera; Tenebrionidae; Coniontini; Eusatti). Occa- sional Papers of the California Academy of Sciences, No. 141, 104 pages, 142 figures, 1984. — Eusattus LeConte and Conisattus Casey constitute a monophyletic subtribe of Coniontini, the Eusatti. The Eusatti are herein considered coordinate with the Conionti (Coelus, Coniontis) and the Branchi (Bronchus, Anectus, Oxinthas). The characters used to define these lineages include internal skeletal features and the configuration of the internal female reproduc- tive tract, as well as characteristics of legs, antennae, and other external structures. Among the Eusatti, Conisattus is most pleisiomorphic, sharing only one of the synapomorphies uniting the species of Eusattus. Eusattus comprises 8 lineages, recognized here as species groups based on configuration of the meten- dosternite, female reproductive tract, legs, antennae, and elytra; a total of 39 species and 13 subspecies are included. New species are: Eusattus arenarius, aridus, catalinensis, catavinus, cedrosensis, ceralboensis, cienegus, ciliatoides, crypticus, franciscanus, hirsutus, minimus, pallidus, phreatophilus, planulus, rudei, and vizcainensis. New subspecies are: E. dubius abditus, dubius arizonensis, dubius setosus, muricatus diabloensis, pallidus adustus, pallidus imma- culatus, pallidus pallidus, and politus cruzensis. Several new synonymies are proposed. Keys are provided to the genera of Coniontini and to the species groups, species, and subspecies of Eusattus. Systematics of Eusattus and Conisattus (Coleoptera; Tenebrionidae; Coniontini; Eusatti) John T. Doyen Introduction The Coniontini comprise one of the major ele- ments of New World Tentyriinae, with approx- imately 200 species recognized in the 1908 monograph by T. L. Casey. Although this figure is somewhat bloated because of Casey's tendency to recognize individual variation nomenclatur- ally, there will probably not be a drastic reduc- tion, as many species are still undescribed. The Coniontini may be divided into 4 groups of closely related genera, as detailed later in this paper. One of these, represented by the genus Coelus, was the subject of a previous revisionary study (Doyen 1976). Eusattus LeConte (sensu Triplehorn 1968) and Conisattus Casey consti- tute the subtribe Eusatti, which is the subject of the present work. The subtribes Conionti and Branchi, defined below, will be addressed in later treatments. Available keys and descriptions preclude re- liable determination of Eusattus. Even with a reference collection, Casey's cumbersome clas- sification makes identification tedious. Conse- quently, two of the objectives of this study are to provide more useful keys and descriptions of adults and to accurately depict distributions. Larvae of several species have been associated, but are relatively conservative and less useful taxonomically; they will be treated separately. Beyond these practical goals, the Eusatti are interesting systematically because of the unusual amount of variation in external features. Several internal structures have proven to be unexpect- ably variable as well, and an attempt has been made to place these patterns of morphological variation into a phylogenetic framework, using a cladistic approach. Finally, patterns of distri- bution in Eusattus are of some biogeographic interest and will be discussed in this study. Acknowledgments This study was facilitated by the cooperation of many people. C. A. Triplehorn, Ohio State University, generously provided specimens which he had been accumulating for many years. Elbert Sleeper, Long Beach State University, invited me to examine his unmounted material from Baja California, which yielded many valuable speci- mens. M. J. D. Brendell, British Museum (Nat- ural History), very kindly sent the type series of the species described by Champion, allowing me to study them at length. The same courtesy was extended by the Burke Museum, University of Washington, for the type of Conisattus nelsoni, and by A. F. Newton, Museum of Comparative Zoology, Harvard University, for several Horn types. Special efforts to collect Eusattus for this study were made by C. E. Griswold, J. K. Liebherr, J. A. Powell, and E. I. Schlinger, University of Cal- ifornia, Berkeley; P. A. Rude, University of Cal- ifornia, Davis; M. E. Irwin, University of Illinois; S. E. Miller, Santa Barbara Museum of Natural History and Harvard University; and J. E. Gil- laspy, Texas A and I University, Kingsville. Early phases of the field work were supported by NSF Grant BMS 74- 1 7924 and by the Scripps Institution of Oceanography, which sponsored the Dolphin Expedition that visited several is- lands in the Gulf of California under the lead- ership of L. Cheng in April and May, 1974. Permission to do field work on San Clemente and San Nicolas islands was granted by the De- partment of the Navy. Pacific Missile Range; per- mission to work at the Zzyzx Springs Station was granted by the California State University Sys- tem. The excellent illustrations of beetles were done by Celeste Greene and Carolyn Mullinex Tib- bets; Ms. Tibbets also made most of the illus- trations of structural features and the charts. Bra- conid parasites were determined by C. C. Loan, of Canada Agriculture. Finally, the work would have been impossible without the cooperation of the following insti- tutions and individuals who loaned the speci- mens under their care: R. Aalbu collection; American Museum of Natural History, New York (L. Herman); Arizona State University, Tempe (F. Hasbrouck); Brigham Young University, [1] OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Provo, Utah (S. Wood); British Museum (Nat- ural History), London (M. J. D. Brendell); Burke Museum, University of Washington, Seattle; California Academy of Sciences, San Francisco (H. Leech and D. Kavanaugh); California De- partment of Food and Agriculture, Sacramento (F. Andrews); California State University, Long Beach (E. Sleeper); Canadian National Collec- tion, Ottawa (D. Bright, J. E. H. Martin, and J. M. Campbell); Cornell University, Ithaca, N.Y. (L. Pechuman); Enns Museum, University of Missouri, Columbia (T. Yonke and E. Riley); Essig Museum of Entomology, University of Cal- ifornia, Berkeley (J. A. Chemsak); Field Museum of Natural History, Chicago (H. Dybas and E. Smith); Kirby W. Brown collection; Los Angeles County Natural History Museum (J. Donohue and R. Snelling); Michigan State University, East Lansing (R. Fischer); Museum of Comparative Zoology, Harvard University, Cambridge, Mass. (J. F. Lawrence and A. Newton); Ohio State Uni- versity, Columbus (C. A. Triplehorn); Oregon State University, Corvallis (P. Ritcher); Peabody Museum, Yale University, New Haven, Conn. (K. Brown); Philadelphia Academy of Sciences (W. Moss and H. Roberts); Battelle Pacific Northwest Laboratories, Richland, Wash. (L. Rogers); Santa Barbara Museum of Natural His- tory, Santa Barbara, Calif. (S. Miller); Texas A and I University, Kingsville (J. Gillaspy); Texas A and M University, College Station (H. Burke and S. Merrit); University of Arizona, Tucson (F. Werner); University of Arkansas, Fayetteville (R. Chenowith); University of California, Davis (R. Schuster); University of California, Irvine (P. Marsh); University of California, Riverside (S. Frommer); University of Idaho, Moscow (W. F. Barr); University of Kansas, Lawrence (P. Ash- lock); U.S. National Museum of Natural History, Washington, D.C. (T. Spilman); Utah State Uni- versity, Logan (W. J. Hanson); Washington State University, Pullman (W. Turner); D. K. Young collection. Materials and Methods Collecting Techniques Most species of Eusattus are retiring, noctur- nal creatures which are not commonly collected except by specialists. The fossorial species (E. muricatus and ciliatus species groups) are seldom encountered on the surface except at night, when they may be found by searching with lanterns. These species may also be screened from sand. For some, such as E. ciliatus, ciliatoides, and pallidus, practically all individuals in collections have been obtained in this manner. The sand beneath low-growing vegetation is usually most productive, and frequently yields larvae as well as adults. Pitfall traps are useful for collecting some species, especially if the substrate is sandy, so that emplacement is easy. Practically any con- tainer is serviceable, but disposable plastic cups, 7-10 cm in diameter, are lightweight and con- venient. Larger traps containing ethylene glycol preservative may be left in the ground for periods up to a year. They are useful for remote areas which cannot be visited frequently. Most spec- imens of £. aridus were trapped in ethylene gly- col pitfalls. Many species of the reticulatus and convexus groups hide in the litter beneath low shrubs, ven- turing forth infrequently. To collect these, it may be necessary to uproot the plants and search through the debris. Nearly all specimens of E. erosus, catavinus, and minimus were found in this manner. Rearing Eusattus specimens are easily transported in containers with a bit of soil or litter. Adults will usually oviposit, and larvae may be reared to intermediate instars in shoebox-size containers of soil or sand, as described in Doyen 1972. As the larvae grow they gradually die, and I have obtained pupae of only a single species. Much of the mortality is apparently due to cannibalism, and individual containers for large larvae would probably be helpful. Dissection and Measurement Dissections of adults were made from dried specimens which were softened in hot water, par- tially dismembered, and then placed in hot KOH to remove soft tissues. Mouthparts were stored on depression slides in glycerine jelly. The cu- ticular parts of the female genital tube were cleaned of tracheae, stained in chlorasol black E, and stored on depression slides in glycerine jelly. The stained tube may be transferred di- rectly to the warmed jelly without distortion, ex- cept for collapsing of the lumen of the sper- mathecal accessory gland. For additional details of preparation, see Tschinkel and Doyen (1980). DOYEN: SYSTEMATICS OF EUSATTUS AND COMSAIITS Measurements of body dimensions were made to the nearest 0.1 mm, using calipers (M. P. J. Gauge and Tool Co., England). Measurement of mouthparts, tarsi, antennal segments, and most other structures was done with an ocular grid and micrometer; these measurements are listed as decimals in the descriptions. The size of some features which were difficult to measure, partic- ularly curved structures, were estimated relative to other body parts; these are described as frac- tions. The body dimensions included with each description were measured as follows: elytral length (EL) is the greatest linear midline distance from the anterior margin of the scutellum to the elytral apex; pronotal length (PL) is measured along the midline; elytral and pronotal widths (EW and PW) are maximum widths; body depth (BD) is the maximum distance between dorsum and venter, normal to a frontal plane through the body. Ratios of antennal and tarsal segment lengths are listed in ocular units, all other mea- surements in metric units. Morphological Terminology Descriptive terminology generally follows Doyen (1966), unless more precise terms are re- quired. The terminology of Tschinkel and Doyen (1980) is used for the internal female reproduc- tive tract (see Figs. 1-12) and ovipositor. Two features are exceptionally variable in Eusattus, and are mentioned repeatedly in the descriptions and keys: 1 . The elytral epipleura may become gradually narrowed toward the apex (Fig. 38), or may be abruptly narrowed just behind the humeri (Fig. 36). In E. robustus, the epipleural ridges mark the lateral extent of the elytra (Figs. 39-40). In all other species, the elytra are inflated so that the epipleura become ventral and more medial in position, at least posteriorly (Figs. 35-38). The surface between the epipleural ridge and the lat- eralmost extent of the elytron is here termed the pseudepipleuron. In several species the elytra are secondarily ridged or carinate, forming a pseud- epipleural ridge (Fig. 36). 2. The spination of the anterior legs is of ex- ceptional taxonomic value. The important de- scriptive terms and orientation used here are il- lustrated in Fig. 4 1 . Treatment of Data To conserve space, collection data are con- densed as follows: For common species, the dis- tribution is summarized and illustrated with a map. For less common species, the collection localities and dates, along with any ecological information, are listed, but collectors and insti- tutions of deposition are omitted. Complete data are provided for type series of all newly described species. Abbreviations of depositories are as fol- lows: BMNH British Museum (Natural History), London CAS California Academy of Sciences, San Francisco CDFA California Department of Food and Ag- riculture, Sacramento CNC Canadian National Collection, Ottawa CSULB California State University, Long Beach EME Essig Museum of Entomology, Univer- sity of California, Berkeley MCZ Museum of Comparative Zoology, Har- vard University, Cambridge, Mass. OSU Ohio State University, Columbus RLA Rolf L. Aalbu collection UCI University of California, Irvine USMNH United States Museum of Natural History, Washington, D.C. Several institutions are worthy of special men- tion because of the richness of their holdings. Foremost is the California Academy of Sciences, which probably has the largest collection of Pa- cific slope Tenebrionidae in existence. This col- lection is of historical significance because of the work of F. E. Blaisdell, and includes cotypes of many of the species G. C. Champion described from Mexico and Central America. All type- specimens designated in this work are deposited in the California Academy of Sciences collection. The collection of the California State Univer- sity at Long Beach is notable for its holdings of material from Baja California, accumulated al- most entirely by Elbert Sleeper and his students over a period of many years. All parts of the peninsula are represented, and for many areas no other material is available. The collection of the California Department of Food and Agriculture includes extensive hold- ings of Coleoptera from the desert regions of western North America, especially from sand dunes. This collection also contains significant holdings from Baja California. OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 Future Work Future taxonomic investigation of the Eusatti would be most productively concentrated in Mexico, including Baja California. Only a few hundred museum specimens of all species com- bined have been accumulated from mainland Mexico. Two species are known from a single locality, and for several the type series described by Champion (1892) are the largest collections in existence. Reconnaissance of arid regions of the tier of states from Nuevo Leon to Veracruz is particularly needed. Significant material from Baja California has been accumulated in recent years, but large areas are still unknown entomologically. About half the species of Eusattus occur only in Baja Cali- fornia. Many of these, recorded from one or a few localities, are newly described herein. Some of the species described nearly a century ago re- main rare in collections, with no extensive series from any locality. The islands in the Gulf of California present a special problem. Early expeditions of the Cal- ifornia Academy of Sciences failed to turn up Eusattus on several islands where they were found by later parties. Moreover, Blaisdell ( 1 943) men- tions specimens from Isla Cedros and Isla Ce- ralbo which cannot now be located in the Acad- emy collection. Probably all the islands need to be revisited during favorable seasons. Biology The biological requirements of Eusatti are very incompletely known. Habitat preferences of in- dividual species are summarized in the taxonom- ic treatment. General features of the life history are discussed below. The only detailed biological information on Conisattus is included in the Rogers et al. ( 1978) survey of the feeding preferences of Tenebrion- idae on the Hanford Test Site in south-central Washington. Gut-content analysis recorded frag- ments of foliage from 12 species of angiosperm plants, as well as arthropod parts, pollen, and seeds. Descurainia pinnata (Walt.) Britton con- stituted 37% of the material consumed, but no other item accounted for more than 14%. Forbs, shrubs, and grasses were among the most fre- quently consumed items, suggesting quite gen- eral feeding preferences. According to Boddy ( 1 957), Conisattus inhab- its sand dunes, and Rogers et al. (1978) report that it occurs as a rare inhabitant of sandy soils. Its morphological features suggest a life on the surface or in upper layers of soil or debris, rather than a strictly fossorial existence. The elongate, relatively slender body is similar to that of sur- face dwellers such as Coniontis rather than the globose bodies of burrowing forms such as Eu- sattus muricatus. The foretibiae are also gener- alized, without the long setal fringes present in strongly fossorial species of Eusattus. The only characteristic indicative of fossorial habits is the well-developed lateral body fimbriation, but some surface-dwelling groups (e.g., many Pedinini) are also fimbriate. Habitat Preference and Phenology Biological knowledge of Eusattus is largely limited to data which can be gleaned from collec- tion labels. More specific information exists for a few western species. Based on mode of life, 2 cohorts of species may be recognized. Members of the reticulatus, convexus, and depressus species groups are primarily dwellers in woodland or grassland habitats. Most of the species in these groups are active as adults during the high-pre- cipitation summer months which characterize the southwestern United States and Mexico. This activity pattern is particularly evident in E. re- ticulatus, which is well represented in collections, but pertains to E. depressus, pons, venosus, mex- icanus, franciscanus, laevis, planulus, and south- ern Baja California populations of costatus as well. E. convexus has been collected in all months of the year in southern New Mexico, but the bulk of the records are from summer and fall. E. min- imus, obliteratus, and nitidipennis are poorly sampled, but the limited information suggests adult activity in the summer and early fall months. Several species in Baja California de- viate from this pattern. Most records for E. ca- tavinus and for E. costatus in northern Baja Cal- ifornia are from late winter to early summer months, when precipitation is concentrated. E. aridus, which is restricted to the Vizcaino region where precipitation is seasonally unpredictable (Hastings and Turner 1 965), appears to be active in the summer, but collections are few. Collec- tions of the remaining species of this cohort (E. catalincnsis, cedrosensis, ceralboensis, secutus) are so few that no pattern is evident. The species discussed above are all surface dwellers which spend the daylight hours hidden DOYEN: SYSTEMATICS OF EUSATTUS AND CONISATTVS in the litter accumulated about the bases of shrubs or trees, or occasionally under stones, boards, or other objects. At night, or infrequently during the day, individuals leave their hiding places to forage, but even then may not venture far from the canopies of shrubs. The distribution of many of these species is highly local, especially in desert regions, where they occur primarily in riparian situations. Such factors have often resulted in meager collections, even though a species may be locally abundant. For example, E. erosus manuelis was common about the bases of Atri- plex growing just behind the beach at Bahia San Gabriel, Isla Espiritu Santo (J. Doyen Lot 74C9), yet at night no beetles were discovered foraging. In this case, nearly all known specimens resulted from this one collection. Similarly, E. catavinus was found to be common beneath shrubs or for- aging nearby in the sandy arroyo bottom, but was entirely absent from the surrounding slopes (J. Doyen Lot 74D1). E. minimus occurred only in the litter about bases of low shrubs growing along a road berm in Nuevo Leon, Mexico (J. Doyen Lot 81F3); grassy or barren areas only a few meters away yielded no beetles. A second cohort of species inhabits desert or semidesert situations, almost always in areas of sandy substrate. The more generalized members of this subgroup are in the product us and robustus species groups and E. dubius of the ciliatus group. These beetles are surface dwellers, sheltering un- der shrubs, stones, or other objects, much as the species discussed above. The more specialized members of this desert-inhabiting cohort are adapted for a fossorial life in aeolian sand. These species, comprising the muricatus, rudei, and cil- iatus species groups, exhibit the most profound morphological modifications of any Eusattus. These adaptations, which involve body shape, setation, and structure of legs and antennae, are discussed below in the section "Cladistic Rela- tionships among Eusattus Species Groups." E. dubius, productus, and difficilis have been collected mostly in the winter and spring months, when local precipitation is concentrated and temperatures are less extreme. Members of the muricatus, ciliatus, rudei, and robustus species groups are active as adults throughout the year, except in the colder parts of the range of muri- catus, where winter activity is curtailed by low temperatures. E. robustus and politus, though surface dwellers, are probably able to remain ac- tive, because island habitats are extremely mar- itime, with significant advective precipitation from summer fog. Some of the other species (e.g., E. pallidus, ciliatus, ciliatoides, crypticus, rudei) occupy maritime situations, but their ability to escape unfavorable conditions by burrowing into the deeper layers of sand is probably more im- portant. For example, both E. pallidus and E. franciscanus occur on Isla San Francisco in the Gulf of California, but pallidus, a fossorial species, occurs through the dry season, while francisca- nus, a surface dweller, is restricted to the wet summer period and the following few months. E. muricatus and dilatatus remain active throughout the year in the Colorado and lower Mojave deserts, despite extremely high summer temperatures (Andrews et al. 1979 and personal observation). The daytime shelters of the fossorial species are usually concentrated in the sand beneath plants which shade the surface. The beetles rest in the sand a few to many centimeters deep, de- pending on the degree of shading. Activity typ- ically begins well after dark, when the surface layer has cooled to a tolerable temperature. On overcast days, when the sand is cooler, activity may begin in the middle of the day. By sunrise most of the beetles have dug in for the day, and by the time the surface temperature begins to rise, all have retreated into the sand. Life History and Feeding Feeding habits have been studied only in E. muricatus (Rogers et al. 1978) and E. convexus (Kumar et al. 1976; Lavigne 1980). Both species consume small quantities of arthropod parts as well as foliage from a wide variety of plants, but concentrate on one or a few species. It seems likely that the favored food must change season- ally and geographically, and these species are best regarded as generalists. Probably the feeding habits of other species of Eusattus are similar. Eggs are deposited in soil or sand, hatching in about 10-14 days in the laboratory. Under lab- oratory conditions, larvae reach large size in 6- 8 months. Larvae of E. reticulatus cultured in August 1971 pupated in late July 1972. A one- year life cycle may be prevalent throughout the genus. All Eusattus and at least some Coniontis pro- duce audible sounds by tapping the abdomen against the substrate (Tschinkel and Doyen 1 976). The tapping rate, the length of tapping pulse OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 and their modulation are characteristic in species which have been studied. Only males tap, and a role in sexual communication seems likely. Predators and Parasites The Channel Island fox (Urocygon) feeds on several tenebrionids, including E. robustus (Doy- en 1974a). Insectivorous birds, reptiles, and oth- er small mammals are known to consume other Tenebrionidae, even those with defensive secre- tions. They undoubtedly include Eusattus in their diets, but published records are unavailable. The most important predators of many Tenebrioni- dae may be invertebrates. Polis (1979) lists E. muricatus as one of the mainstays (13.5%) of the diet of the scorpion, Paruroctonus mesaensis, and Tenebrionidae constituted 42.3% of the total food items consumed. Black widow spiders common- ly kill tenebrionids, including large species with defensive secretions. Judging from webbed corpses I have observed, they are important predators of E. robustus, reticulatus, and cata- vinus, as well as ofcrypticus (E. I. Schlinger, pers. comm.). Unidentified spiders are predators ofE. ciliatoides, dubius, minimus, pallidus, rudei, and vizcainensis. There is a single instance of ant- lion (Myrmeleontidae: Neuroptera) predation on an adult E. dilatatus (J. Doyen Lot 72D2). Pred- ators of larval Eusattus are not recorded in na- ture, but in laboratory situations myrmeleontid and therevid (Diptera) larvae voraciously con- sume tenebrionid larvae, including those of Eu- sattus. It is likely that Therevidae are specialists on Tenebrionidae (M. E. Irwin, pers. comm.). Adults of many genera of Tenebrionidae are attacked by parasitoid wasps of the genus Mi- croctonus. I have reared M. eleodis (Viereck) from E. cienegus, dilatatus, minimus, and muricatus. The parasitoids may be abundant: among a col- lection of 89 E. dilatatus taken on 27 March 1972, 22 individuals were infested. The wasp larvae live in the thoracic and abdominal hae- mocoel, causing no apparent symptoms until shortly before their emergence from the beetle. One to several parasitoids may mature in a single beetle; in the similar-sized Ed rotes ventricosus LeConte, up to 30 larvae were dissected from a single host. The mature larvae emerge from the anal end of the host, falling to the substrate. A few hours before emergence, infested beetles be- come moribund and unable to use their hind legs in walking. Parasitism invariably results in death of the host in a few hours to about a day after emergence. After approximately an hour of wan- dering, the larvae spin oval cocoons incorporat- ing particles from the substrate. Adults emerge in the laboratory after 15-21 days, and within 1-2 days begin seeking hosts. They run along behind the beetles, attempting to insert the ovi- positor between the elytra and 7th sternite when- ever the beetles stop moving. Microctonus eleodis apparently attacks diverse genera of Tenebrionidae. In the laboratory, wasps reared from a single collection of Edrotes ven- tricosus successfully parasitized Edrotes arens La Rivers, Eleodes gracilis LeConte, Eleodes bea- meri Blaisdell, Eleodes extricata (Say), and Eu- sattus dilatatus, but failed to parasitize Crypto- glossa verrucosa LeConte and C. laevis LeConte. The broad host range probably compensates for the absence of some hosts (such as Eusattus min- imus) during much of the year. Higher Classification of Coniontini The tribes Coniontini and Branchini were es- tablished by Lacordaire (1859) and LeConte (1862) respectively. The Coniontini were com- prehensively treated by Casey (1908), who rec- ognized about twice the number of genera exist- ing earlier. With the exception of Conisattus, these new taxa resulted from splitting previously es- tablished genera. Eusattus was especially affect- ed, being divided into 6 genera. Triplehorn (1968) reunited most of the Eusatti under the old genus Eusattus, based on intermediacy in the differ- entiating external characters used by Casey. In particular, Triplehorn noted, as had LeConte (1866), that the configuration of the epipleuron in Eusattus could not be divided into discrete character states, because of transitional condi- tions between the extremes. My preliminary analysis of higher classifica- tion of the Coniontini (Doyen 1972) supported Triplehorn's consolidation of the Eusatti and placed the remainder of Casey's ( 1 908) genera in synonymy under Coniontis Eschscholtz and Coe- lus Eschscholtz. At the same time, the 3 genera of the tribe Branchini were transferred to the Coniontini. These changes were based on com- parison of mouthpart, genital, and internal skel- etal structures, as well as external features of all the genera except Conisattus Casey and Anectus Horn. The relationships of Conisattus, which have proven problematic, will be discussed at length below. The present analysis stems from an attempt to DOYEN: SYSTEMATICS OF EUSATTUS AND COMS 1/77 S understand the cladistic structure of the genus Eusattus. In order to estimate character-state po- larities, all genera of Coniontini (including Bran- ching were examined. For some characters the direction of change remains unclear; correct po- larities can only be determined by looking at a spectrum of related tribes. Since many of these are large and taxonomically diverse themselves, such an undertaking is beyond the scope of this analysis. It should be noted that only adult fea- tures are considered here, since larvae are in- adequately characterized. Nevertheless, even though preliminary, an estimate of cladistic re- lationships among Coniontini is now possible. Character States Characters and character states are discussed below. Character-state distributions are shown in Fig. 13, numbered as follows. 1. Epistomal Shape.— An emarginate episto- mum probably evolved independently many times in different groups of Tentyriinae, but is probably synapomorphous in Coniontini. The primitive, entire epistomum is widespread in many tribes. 2. Antennal Length. — Antennae in most Coniontini and other tribes of Tentyriinae are about as long as the prothorax. This is the prim- itive condition. In taxa which burrow through loose sand, the antennae often become much shortened, sometimes with the fusion of seg- ments. Shortened antennae are present in all species of Coelus and in Eusattus dilatatus, also a burrower in aeolian sand. The antennae of E. dilatatus are moniliform, with no indication of a club, and differ in proportional segment lengths from those of Coelus. These and other differences indicate convergence in antennal form. 3. Antennal Configuration. — Filiform or setaceous antennae are widespread in Tentyri- inae. In most Coniontini the antennae are grad- ually and slightly enlarged toward the apex. In Coelus and a few Eusattus, the terminal 3 seg- ments form a very weak club (Fig. 34); for the analysis here, these conditions are included in the primitive state. In Branchus, Anectus, and Oxinthas, the 3 terminal segments form a dis- tinct and abrupt club (Fig. 27), which is here considered derived. 4. Mentum Size.— It is unclear whether a small mentum is primitive to Coniontini, or derived. The small mentum is nearly ubiquitous in Te- nebrioninae, Lagriinae, and related families of Heteromera, where it is undoubtedly primitive. A large mentum occurs in several winged ten- tyriine tribes (Tentyriini, Eurymetopini, Epitra- gini, Evaniosomini), as well as many wingless, obviously derived tribes. A small submentum, though uncommon, occurs in some winged Ten- tyriinae, such as Vacronini (sensu Doyen and Lawrence 1979), as well as in miscellaneous tribes such as Anepsiini, Cnemoplatiini, Lachnogyini, Stenosini, and Coniontini. The first distribution suggests that the large mentum may be primitive to the entire Tentyriinae, with several secondary reductions, as assumed here. Alternatively, the enlarged mentum could be multiply derived. Fi- nally, the division into taxa with large versus small mentum could represent a primary di- chotomy, but this is not obviously supported by other characters. 5. Development of Submentum.— The an- terior region of the gula is delimited by sutures to form a large, distinct submentum in many tribes of Tentyriinae (Tentyriini, Eurymetopini, Epitragini, Asidini, etc.), suggesting that this con- dition is primitive. In most Coniontini the sub- mentum is reduced to a small, transverse sclerite (Fig. 28). In Eusattus cienegus and in Branchus, Anectus, and Oxinthas, the submentum is further reduced and not visible externally (Fig. 29; Doy- en 1972, fig. 24). 6. Gular Configuration. — Primitively, the gular sutures are separate their entire length, con- verging anteriorly. This condition is present in most Coniontini. In Eusattus cienegas, the su- tures are anteriorly confluent or nearly so. In Branchus, Anectus, and Oxinthas, the sutures are confluent in their apical third. Confluent gular sutures are 100% correlated with loss of the sub- mentum in Coniontini. 7-8. Foretibia Shape. — The primitive config- uration in Tentyriinae is subcylindrical or slight- ly flattened apically. In sand-burrowing or sand- swimming forms, the foretibiae are commonly specialized as lamellate digging tools, often of highly idiosyncratic configuration. In Coniontis and Coelus, the foretibiae are not significantly modified from the primitive condition. In Bran- chus, Anectus, and Oxinthas, the apex is pro- duced as a short hooklike projection (Fig. 24). In Conisattus and Eusattus, the tibia is strongly flattened and gradually expanded apically as an attenuate, acute process (Fig. 41). The numerous differences between the Eusatti and Branchi in- dicate that these states developed independently. OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 9-10. Protarsal Shape. — In Coelus, the basal protarsomere is uniquely modified as a large, la- mellate digging structure (Doyen 1972, fig. 20). In all other Tentyriinae this tarsomere is short and subcylindrical. The basal protarsomere length (not including processes) is subequal to that of the 2nd tarsomere in most Coniontini and other Tentyriinae. In Eusattus, the basal tarsomere is subequal to the next 3 tarsomeres combined. 11-12. Pronotal Setal Fringe.— Typically in Tentyriinae the hypomeral region is glabrous or nearly so, and this is the condition in most genera of Coniontini. Burrowing Tentyriinae often have fimbriate lateral body margins. Usu- ally the setae are set on the carina, as in Coelus and Conisattus (Fig. 25). Such setal fringes are probably developed from short carinal setae such as those present in Eusattus muricatus and in Branchus. In Eusattus, setae inserted on the hy- pomeron well below the carina form a prominent pronotal fringe (Fig. 26). This state is uncommon in Tentyriinae, and probably developed inde- pendent of carinal setae. 13. Epipleural Shape. — In most Tentyriinae the epipleuron comprises a relatively narrow strip bordering the elytron laterally. In Coniontini, as in many other Tenebrionidae, the epipleural re- gion faces ventrad because of inflation of the hind body (Figs. 35-40). In the reticulatus species group of Eusattus and in the Branchi, the epipleuron is narrow posteriorly but very broad anteriorly, where it coincides with the lateral elytral margin (Fig. 36). In other species groups of Eusattus, epipleural configuration is variable, and the spe- cialized, broadened epipleura apparently evolved independently in the Branchi. 14. Setal Shape. — Setae in Tenebrionidae are characteristically hairlike, which is the primitive state. In the Branchi, the dorsal body setae are short and clavate; this condition is especially ev- ident on the elytra. In Eusattus puberulus, the dorsal setae are squamiform and superficially similar to those of the Branchi. 1 5. Metendosternite Configuration. — The primitive arrangement in Coniontini, as in other Coleoptera, is a Y-shaped metendosternite, with the apices of the arms attached by muscles to the metatergum. In all Eusattus the arms are elon- gate, with the apices connected to the lateral edge of the mesotergum by a very short tendon (Doy- en 1972, fig. 12). This modification has occurred independently in many distantly related tribes of Tenebrionidae, but appears to be a valid syn- apomorphy in Eusattus. 16. Spermathecal Tube Number. — Multiple (6-8) spermathecal tubes are widespread in Te- nebrionidae, especially in Tentyriinae, and ap- parently represent the primitive condition (Tschinkel and Doyen 1981). Reduced numbers of tubes are characteristic of various genera of Coniontini (Figs. 1-6), but it is impossible to place these different states into a simple trans- formation series without disrupting other syn- apomorphies. Moreover, spermathecal tubes are sometimes branched, and occasionally vary in number within a single species. Interpretation of polarity in this character is further complicated by differences in the shape of the tubes, described below. For the purposes of this analysis, each tube number is assumed to have arisen indepen- dently by reduction from the primitive condi- tion. 17. Spermathecal Tube Symmetry. — In most Tentyriinae the multiple spermathecal tubes are similar in size and shape. In Coniontini one of the tubes is reduced in size and sometimes at- tached by a portion differentiated as a duct (Figs. 1-6) (in Coelus and Coniontis punctulata Horn, there is only a single tube). As far as is known, this configuration is unique among Tentyriinae, but spermathecal arrangement is not yet ade- quately surveyed. 18. Spermathecal Tube Shape. — The prim- itive condition, widespread in Tentyriinae, is long, slender tubes, usually tightly coiled (Fig. 1). In Coniontis punctulata and C. lata LeConte, the tube is somewhat shorter and thicker (Figs. 3- 4), and in other Coniontis the tubes are very short and thick (Fig. 2). In Eusattus the tubes may be long, slender, and tightly coiled or short, thick, and irregularly contorted. Because of the other synapomorphies of Eusattus, it seems very likely that shortening of the spermathecae has inde- pendently arisen several times in Coniontini. Ac- cordingly, no synapomorphies for character 18 are shown in Fig. 13. 19. Spermathecal Accessory Gland Duct Shape. — Typically, this duct is long and thin in Tenebrionidae, including Coniontini (Figs. 1-2 and 6). In Coelus, the duct is relatively thick and short (Fig. 5), and in Coniontis (=Coelotaxis) punctulata it is intermediate (Fig. 3). 20. Cuticular Encrustations. — Most Ten- tyriinae have glabrous or subglabrous cuticle— or if setae are present, the cuticle itself is bare. In Branchus, Anectus, and some Eusattus of the reticulatus group, the cuticle, especially of the elytra, is encrusted with soil particles. The en- DOYEN: SYSTEMATICA OF Ei'SATTL'S AND ( O.XISA III S •SD Figures 1-6. Cuticular parts of female genital tracts. B = bursa; SAG = spermathecal accessory gland (stippled); SD = spermathecal duct(s). 1. Conisattus rectus: a. entire tract; b. spermathecal ducts, enlarged. 2. Coniontis (Coniontis) pectoralis Casey. 3. Coniontis (Coelotaxis) punctulata. 4. Coniontis (Comontides) lata: a. entire tract; b. spermathecal ducts, enlarged. 5. Coelus globosus LeConte. 6. Branchus obscurus Horn. crusting habit is derived independently in some other tentyriine tribes such as Cryptoglossini, where the material is a waxy secretion (Hadley 1979), and is present in many other families of Coleoptera (Lawrence and Hlavac 1979). Prob- ably the double occurrence in Coniontini is con- vergent. Cladistic Relationships The genera of Coniontini comprise 2 groups, the Branchi and the combined Conionti and Eu- satti (Fig. 13). Branchus, Anectus, and Oxinthas (subtribe Branchi) share synapomorphies in sub- mentum and gular configuration, antennal shape, elytral vestiture, and epipleural shape. The last 10 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 SAG Figures 7-9. Cuticular parts of female genital tracts. 7. Eusattus reticulatus: a. entire tract; b. c. d. spermathecae from 3 individuals, showing variation. 8. E. productus: a. entire tract; b. spermathecal ducts, enlarged. 9. E. depressus: a. entire tract; b. base of spermathecal ducts. also occurs in the reticulatus group of Eusattus, probably independently. The Branchi are highly derived compared to their sister group, the com- bined Conionti and Eusatti, but show little dif- ferentiation among genera. Ancctus is virtually identical to Bronchus in important characters. and should be placed in synonymy. These taxa are superficially similar to members of the reticu- latus species group of Eusattus. Oxinthas is su- perficially similar to Coniontis. The remaining genera (Conionti plus Eusatti) share no convincing synapomorphies, but are DOYEN: SYSTEMATICS OF EUSATTUS AND ( ON1S. 1 l/l S 11 Figures 10-12. Cuticular parts of female genital tracts. 10. Eusattus crypticus: a. entire tract; b. spermathecae, enlarged. 1 1. E. robustus: a. entire tract; b. spermathecae, enlarged. 12. E. ciliatus. united by a high level of overall similarity, es- pecially among larvae, and synapomorphies may eventually become apparent when larvae of re- lated tribes are characterized. The Eusatti universally share a single character which is unquestionably derived. This is the la- mellate shape of the protibiae. In addition, Co- nisattus has 4 long, slender spermathecal tubes, a feature believed primitive but found elsewhere only in some Eusattus, and quite different from the spermathecal arrangement in other Conion- tini. In most other characters Conisattus is plei- siomorphic and superficially resembles Conion- tis. Casey (1908:146) recognized this similarity, and on this basis I originally placed Conisattus in synonymy under Coniont is (Doyen 1972); but on the basis of the single synapomorphy I later (Doyen 1977) removed Conisattus to Eusattus. Its pleisiomorphic internal skeletal anatomy, to- gether with unique apomorphies such as the pro- notal setation, make it appropriate to reassign Conisattus to generic status. Eusattus is distinguished by 3 apomorphies (fusion of notum with metendosternite; hypo- meral setal fringe; protarsal structure) which do not occur elsewhere in Coniontini. In addition, the pronotum in Eusattus usually has a charac- teristic shape, with bisinuate base and produced posterior angles, but a similar configuration oc- curs in Branchus and some Coniontis. In several other important characters Eusattus is variable, sharing apomorphic states with other genera or subtribes. Several of these features are deemed important in the infrageneric relationships of Eusattus, as discussed in the next section. According to the characters included here, the Conionti consist of 2 subgroups. Coelus and Coniontis punctulata (=Coelotaxis Casey 1908) have only a single spermathecal tube, which is relatively shorter than those of Conisattus but longer than those of Coniontis (sensu stricto). Coelus, which is strongly adapted to a fossorial life, is highly derived in a number of other fea- tures. Coniontis, including C. lata (=Conion- tides Casey 1908) has 3-5 spermathecal tubes. In most Coniontis these are very short and thick; in C. lata they are intermediate in length, as in C punctulata. It should be pointed out that no important synapomorphies are shared by the two subgroups of Conionti. They are combined here because of the striking overall similarity between Coniontis 12 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 C0NI0NTI CO CO LU X Q < 1- 1— ^ o o LU «—; z EUSATTI BRANCHI o o CO CO o o , ^ , CO CO CO CO i co co ATTU TUS 3 => co H P 1- x x Z> CO z z z INT ANC \- O Z> _l O o o <* 5 LU LU z 2 ^ 2 CO X DC z o o o o o => o ao < o o o o O LU (16) SPERMATHECAL TUBES = 3 (16) SPERMATHECAL TUBES = 5 (19) ACCESSORY GLAND SHORT Q (2) ANTENNA SHORT (9) PROTARSUS SPATULATE (16) SPERMATHECAL TUBES = I (11) PR0N0TAL CARINA FIMBRIATE (15) METENDOSTERNITE / NOTAL FUSION (10) BASAL PR0TARS0MERE LONG (12) HYP0MER0N FIMBRIATE (16) SPERMATHECAL TUBES = 4 (8) FORETIBIA LAMELLATE (20) ELYTRA ENCRUSTED (7) FORETIBIA HOOKED APICALLY (3) ANTENNA CLUBBED (14) SETAE CLAVATE (13) EPIPLEURAL SHAPE (16) SPERMATHECAL TUBES = 2 (5,6) SUBMENTUM, GULA REDUCED (I) EPISTOMUM EMARGINATE (4) MENTUM SMALL (17) SPERMATHECAE DIFFERENTIATED Figure 13. Cladogram of possible relationships among genera of Coniontini. Numbers explained in text. Closed circles indicate universal presence of derived character states on right; partially closed circles indicate the approximate proportions (numbers of species) when both primitive and derived states are present; open circles indicate absence of information or inapplicability of character. DOYEN: SYSTEMATICS OF EUSATTUS AND CONISATTUS 13 and Coelotaxis, which are scarcely distinguish- able in external features (Doyen 1972:370). If the foregoing analysis is correct, it would be improper on strict cladistic grounds to recognize Coelus at the generic level without also recog- nizing Coelotaxis. This would be inconvenient from a phenetic standpoint, and would unduly complicate generic keys. In the classification adopted here, Coniontis constitutes a monophy- letic clade, in the sense of being convex (Esta- brook 1978). Duncan (1980) and Doyen and Tschinkel (1982) discuss the advantages of this criterion in transforming cladistic results into classifications. Cladistic Relations Among Eusattus Species Groups The extensive morphological variation among species included in Eusattus suggests that their relationships might be well depicted by cladistic analysis. The evident variation includes differ- ences in internal skeletal anatomy and in the configuration of the internal female reproductive tract, as well as numerous differences in leg struc- ture, body proportions, and cuticular sculpturing and setation. Characters were examined over all species of Eusattus and, to estimate polarity, over all gen- era of Coniontini. After preliminary cladistic and phenetic comparisons, 6 1 characters (Appendix A) were selected for the analyses presented here. All characters were binarily coded. Where char- acter-state variation was too complex for simple binary coding, several characters were recog- nized, with additive binary coding (Sneath and Sokal 1973); e.g., see characters 8 and 9 or 18 and 19. After preliminary analyses, it became apparent that several groups of very similar species could be discerned phenetically. In most cases these same groups were distinct cladistically, and are recognized here as species groups. Difficulties in defining species groups and in interpreting ho- mology or character polarity are discussed where appropriate below. Character distributions across all species are listed in Appendix B. One possible cladistic arrangement of the species groups is shown in Fig. 14. Two distinct clusters are evident. The convexus, reticulatus, and muricatus groups constitute cluster A. Their strongest synapomorphy is spermathecal config- uration (Fig. 7)— this highly convoluted, irregu- lar-shaped, and branched structure does not oc- cur elsewhere in Coniontini. Superficially similar spermathecae occur in Coniontis, but in this case the 3-5 or more shortened spermathecal tubes are distinct, with one tube differentiated and set on a short duct (Fig. 2). The shortened tubes in Eusattus are irregular from individual to indi- vidual, and it is usually impossible to determine their number. The differentiated tube, as in other Eusattus, never has a separate duct. The other synapomorphies restricted to cluster A are external features, and might more easily be construed to have converged for adaptive rea- sons. Antennae in these groups are uniformly subglabrous, but antennal pubescence in the clus- ter B taxa is variable. In the ciliatus group the antennomeres are uniformly set with long pu- bescence, while in the difficilis and depressus groups, the pubescence is much shorter and somewhat sparser. This intermediate condition, which occurs in many Tentyriinae, is here con- sidered the primitive character state (Appendix A, chars. 8 and 9). An alternative would be to recognize a single (primitive) state of short, sparse setation. Elytral rugosity is obviously a highly adaptive feature, since it affects sheen and brightness of the dorsum, probably important in camouflage and in heat balance in diurnally active species. In fossorial forms, elytral surface texture is prob- ably important in reducing friction against the substrate. Two other synapomorphies universal to clus- ter A appear in a few cluster B groups: 1 . The subtrapezoidal, asymmetrical, and api- cally angulate terminal antennal segment (Fig. 48), of unknown functional significance, occurs in the rudei and difficilis groups, apparently as convergences. This and other convergent simi- larities are discussed in greater detail below. 2. A relatively short paraproct (Figs. 15 and 16), common in cluster A, occurs in E. crypticus {rudei group) and E. arenarius {ciliatus group). Paraproct length relative to coxite length is a measure of ovipositor length, since the coxite size is relatively constant in terms of body size. Ovi- positor length is probably determined by sub- strate characteristics. In general, the groups with short paraprocts inhabit woodland, grassland, or riparian situations where soils are not particu- larly friable. These beetles are mostly active dur- ing periods of summer precipitation, and eggs 14 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 CLUSTER B CLUSTER A ELYTRA SEXUALLY DIMORPHIC PROSTERNUM ELONGATE INTERCOXAL PROCESS BROAD METENDOSTERNITE /COXAL FUSION SPERMATHECAL TUBES BRANCHED BURSA COPULATR IX PRESENT TERMINAL ANTENNOMERE SHORT LAT. EPIST. EMARGINATIONS DEEP ANTENNA CLUBBED EPISTOMAL SUTURE INCISED FORETIBIAL SPINES CONTIGUOUS ANTENNAE PUBESCENT BODY SUBGLOBULAR INNER FORETIBIAL SETAE LONG PRONOTAL PUBESCENCE MESOSTERNAL SETAE LONG PROSTERNAL SETAE LONG ELYTRA TUBERCULATE FEMORA FIMBRIATE HYPOMERAL FRINGE LONG FORETIBIAL SPINATION PRONOTUM TUBERCULATE PRONOTAL CARINA SETOSE EPIPLEURON ABRUPTLY NARROWED ANTERIOR TIBIAL SPINATION COXITE LONG FORETIBIAL SPINE ROW SHORT TERMINAL ANTENNOMERE ASYMMETRICAL ELYTRA RUGOSE ANTENNA SUBGLABROUS SPERMATHECAE SHORT, CONVOLUTED Figure 14. Cladogram of possible relationships among species groups of EusattUS. Numbers explained in Appendix A; conventions as in Fig. 1 3. DOYEN: SYSTEMATICS OF Ei'SATTi'S AND CONISATTl S 15 are probably less subject to dehydration. The groups of Eusattus with long paraprocts mostly inhabit sandy, desert substrates, where it may be important to deposit the eggs more deeply. How- ever, in other coniontine genera such a correla- tion is not evident, since most inhabit woodlands or grasslands, yet have relatively long paraprocts. Thus it is difficult to assess the importance of ovipositor length as a taxonomic character. Within cluster A, the muricatus group is most distinctive. The members of this group, which are all fossorial, include the most highly spe- cialized Coniontini. However, only 2 apomor- phies are restricted to the muricatus group; both are of unknown function: 1 . Pronotal tubercu- lations, covering the peripheral portion of the disk in the muricatus group, do not otherwise occur in Coniontini. In general, tuberculation and the short, retrorse setae which often accompany it indicate fossorial habits. 2. A fringe of setae on the pronotal carina also occurs in Branchus and Anectus. As in the muricatus group, the setae are sparse and very short. In Coelus and Coni- sattus, the carinal setae are very long, forming a lateral fringe. It is unclear whether the two con- ditions are homologous. The other apomorphies displayed by the muri- catus group are shared with various members of cluster B, especially the ciliatus group. With two exceptions (chars. 2 1 and 4 1 ), all these characters represent adaptations to fossorial life in aeolian sand. Members of the muricatus and ciliatus species groups spend most of their lives within the substrate, and exemplify many of the mor- phological modifications described by Koch (1961) for ultrapsammophilous or "sand-swim- ming" tenebrionids. Among more marked mod- ifications are: 1. the globular, sometimes nearly spheroidal body (char. 51); 2. flattened, paddle- like foretibiae, with the surface area augmented by stiff, projecting setae (char. 40); 3. dense fringes of long setae at the main body articulations, and muffs of setae about the coxal articulations (chars. 33 and 34)— these setae prevent sand grains from entering the joints; 4. short, stiff, retrorse setae on the elytra or other body surfaces (chars. 26 and 27) — these probably assist forward move- ment through the substrate. In the most highly modified species, such as E. dilatatus, further specializations involve shortening of the anten- nae, flattening of an additional pair of legs, or development of setal fringes on them. - COXITE -PARAPRO Figures 15-16. Ovipositors, ventral aspect. 15. Eusattus crypticus, with relatively long coxites. 16. E. costatus, with relatively short coxites. Thus it seems likely that the numerous simi- larities between the muricatus and ciliatus groups result from a remarkable convergence for a com- mon mode of life. Indeed, without the difference in spermathecal structure, the most reasonable classification would combine the two, as was done by Casey (1908). The two remaining groups in cluster A show no convincing synapomorphies. They are com- bined because of their high degree of overall sim- ilarity, especially between E. convexus and E. reticulatus. All members of the reticulatus group have the epipleura suddenly broadened just be- fore the humeri, although in E. araneosus less abruptly so. In this feature, araneosus ap- proaches convexus. The significance of this epi- pleural shape— which also occurs in the Branchi, as mentioned earlier— is unknown. The composition of cluster B is more complex, and the cladistic relationships more problematic. No single synapomorphy unites the entire clus- ter, but several derived features are shared by most members. The members of the difficilis, rudei, ciliatus, and robust us groups all inhabit sandy substrates. With the exception of E. dubius and E. difficilis, the first three groups are restrict- ed to aeolian dunes. Thus it might be expected that several of the synapomorphies would in- volve obviously adaptive features (chars. 1 8 and 35), which are absent in the only group (depres- sus) in this cluster that does not occupy sandy substrates. The significance of a glabrous inner foretibial surface (char. 4 1 ) and tuberculate elytra (char. 21) are unclear, but both features recur in the sand-adapted muricatus group. Within cluster B, most of the species groups 16 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 are distinguished by one or more apomorphies, but synapomorphies uniting groups are uncom- mon. The robustus and rudei groups are united by an unusual sexual dimorphism (char. 25), present in one species of each group. In the fe- males the elytral tubercles become much coarser and closer on the declivity, and in robustus the cuticle becomes shiny. This character does not occur in other Coniontini, nor among other Ten- tyriinae known to me. The depressus and difficilis groups share 2 de- rived characters: 1 . The pair of spermathecal tubes is branched at the base (char. 60), producing the usual number of 4 distal tubes (Fig. 9). 2. The metendosternite arms are held closely against the mesocoxal inflections by a sheetlike tendon, ef- fectively fusing the two surfaces (char. 54). Adhe- sion of the metendosternite and mesocoxal in- flections occurs sporadically in various tribes of flightless Tentyriinae, providing greater rigidity to the body shell. In Eusattus this feature is shared with the muricatus species group. Since the muri- catus group shares no other apomorphies with the depressus-difficilis groups, the metendoster- nite-coxal adhesion has apparently been derived twice. The depressus and ciliatus species groups are most highly derived. The former shows one apo- morphy, an elongate presternum (char. 1 7), which does not occur elsewhere in Coniontini. This pro- longation enables the mouthparts to be almost entirely concealed by the prothoracic fossa when the head is retracted, which probably provides protection against predators. The apomorphies of the ciliatus group are very largely the same as those of the muricatus group, and reflect the strongly fossorial habits. Other apomorphies, such as the weakly clubbed anten- nae (char. 10) and relatively deep, incised epi- stomal suture, (char. 1), are of unknown signif- icance. Relationships Within Eusattus Species Groups Several of the species groups (depressus, dif- ficilis, rudei, and robustus) contain only a pair of species each. The salient differences between members of these pairs are listed in the species diagnoses, and they are not discussed further here. In most of the remaining groups, only a relatively few convincing apomorphies have been discov- ered, or conflicting synapomorphies suggest competing classifications. Consequently, most of the cladograms presented below are incomplete- ly resolved. Reticulatus Species Group This group includes about half the species in Eusattus, and is the most refractory in terms of cladistic reconstruction (Fig. 17). The most dis- tinct species is E. laevis, which shares only the characteristic epipleural shape (char. 20, Fig. 14) with the rest of the group. It is unusual in being almost glabrous, with smooth, shining cuticle; in the latter character it is similar to the species of cluster B. The remaining species share only a single syn- apomorphy, the angulate pseudepipleuron (char. 29). This is a simple feature, but does not occur elsewhere in Eusattus, although it is present in Branchus. Two major clusters are evident. In the costatus subgroup, the ovipositor is relatively long (Fig. 16), the elytra are costate (or secondarily(?) noncostate), and the body tends to be relatively setose. E. cedrosensis, known from a few frag- mentary specimens, is placed here, based on its costate elytra. The reticulatus subgroup, containing the bulk of the species, is not united by apomorphies. Most of the species have reticulate elytra, but there is considerable interspecific variation, and the different textures may well have arisen in- dependently from simple, punctate, or rugose conditions. Within the reticulatus subgroup, sev- eral closely related clusters of species are appar- ent, but their interrelationships are unclear: 1. E. mexicanus, venosus, and pons.— These species share a characteristic body shape (chars. 13 and 50) and cristate pseudepipleural margin (char. 31); the latter feature also appears in E. catalinensis and ceralboensis. These taxa, en- demic to islands in the Gulf of California, are phenetically very similar to E. erosus, which in- habits the Baja California peninsula. The 3 mem- bers of this cluster share their habit of accreting soil particles (char. 28) in the elytral depressions with E. reticulatus and cienegus. However, E. pons, catavinus, and erosus share a high degree of overall similarity, including the very coarsely eroded elytra (char. 23) here considered a derived feature. The confusing synapomorphies of these species are discussed further under "distribu- tional patterns." 2. E. catavinus and erosus. — These are ob- viously sister taxa, based on extremely close overall similarity. Except for their markedly dif- DOYEN: SYSTEMATICS OF EUSATTUS AND CONISATTUS 17 CO co CO Zl CO CO CO CO CO CO CO CO -P' UJ 2 < •> Z) CO 3 3 Z) LU o UJ CO CO _) 2 (/) (C\ 1- _J CO m p. o CO 2 < _J Q Z> o or _i < Lj o 2: UJ ;z 3 CO > o o CO CO < O or < < O X 2 2 tr o _J UJ UJ 5 or LT or 5 UJ UJ o < o Cl CJ o o u_ Q_ CO Z) h- co < Z) —I CD Z> UJ O UJ () C) () () :: :::: o o o o o CO > UJ (20) EPIPLEURON GRAD. NARROWED (23) ELYTRA ERODED (32) PSEUDEPIPL. CRISTATE POSTER. (3D PSEUDEPIPL. CRISTATE ANTER. ( 3 ) MENTUM BROAD ( 5 ) SUBMENTUM SUPPRESSED (30) PSEUDEPIPLEURON SHARP (13) PR0N0TUM TUMID (50) BODY FORESHORTENED (28) ELYTRA ENCRUSTED (22) ELYTRA RETICULATE (57) PARAMERE ATTENUATE (41) ANTERIOR TIBIA SPINOSE (35) FEMORA FIMBRIATE (18) HYPOMERAL FRINGE LONG (34) MESOSTERNAL SETAE LONG (56) COXITE SHORT (24) ELYTRA COSTATE (29) PSEUDEPIPLEURON ANGULATE (19) HYPOMERON GLABROUS Figure 17. Cladogram of possible relationships among species in the E. reticulatus species group. Numbers explained in Appendix A; conventions as in Fig. 13. ferent habitats (see taxonomic account), I would have treated them as subspecies. 3. E. franciscanus, ceralboensis, and catali- nensis. — These species are phenetically similar to E. erosus, but do not share convincing syn- apomorphies with it. They most likely diverged when they were separated on islands in the Gulf of California, probably in the very recent geo- logical past (but see discussion of distributional patterns). 4. E. araneosus.— The relationships of E. ar- aneosus, known from only a few specimens and unavailable for dissection, are uncertain. Its epi- pleural configuration is similar to that of the con- vexus species groups, but it is very similar to E. franciscanus and erosus in other features. Convexus Species Group The species of this group are mostly similar, except for differences in cuticular sculpturing and setation. Consequently it is difficult to specify cladistic relationships (Fig. 18). The three Mex- ican species share relatively smooth, finely sculpted cuticles (chars. 22 and 24), which are 18 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 co CO Z> h- < or LU CO CO LlI Q_ z> X LU > ^ h- Q ^ _l H ^ QQ o ^ O 2 o SJJ (51) BODY SUBGLOBULAR (35) FEMORAL SETAE LONG (18) HYPOMERAL FRINGE LONG (33) PROSTERNAL SETAE LONG (37) FORETIBIA DENTATE (50) BODY FORESHORTENED (22,24) ELYTRA NONRUGULOSE, NONCOSTATE Figure 18. Cladogram of possible relationships among species in the E. convexus species group. Numbers explained in Appendix A; conventions as in Fig. 13. considered reversals and derived within the con- vexus group. These must be considered weak syn- apomorphies at best, since sculpturing varies from strongly reticulate to almost smooth in convexus, sometimes over insignificant geographic dis- tances. The clearest relationship is between E. minimus and E. obliteratus. The characteristic dentate foretibia (Figs. 65-66) is the most con- vincing synapomorphy. The specializations of minimus are similar to those of fossorial species, and the phenetic similarity to E. puberulus (mu- ricatus species group) is striking. The habitat of minimus is the upper layers of soil and litter beneath low shrubs. Muricatus Species Group Two clusters are apparent (Fig. 19). E. phrea- tophilus and E. hirsutus have less globose bodies and a different foretibial spine arrangement (chars. 37 and 40) than those in the E. muricatus subgroup. However, note that E. muricatus dia- bloensis, from northern Baja California, has the spine arrangement of the E. phreatophilus subgroup, which inhabits small sand deposits about the bases of shrubs in alkaline areas. Though unknown to me in nature, E. hirsutus apparently occurs in similar situations, judging from collection records. Members of the muricatus subgroup without exception inhabit aeolian sand, and most of their apomorphies are adaptations to that specialized mode of life. E. muricatus and dilatatus are here placed as sister taxa, but that arrangement must be regarded as uncertain, since it is based on a single feature which varies in muricatus. Ciliatus Species Group Aside from the placement of E. dubius, which is problematic, the cladistic relationships (Fig. 20) are clearer in this group than in any other. The position of dubius is uncertain because it lacks the apomorphies which characterize the rest of the group; it is included in the ciliatus species group because of the high phenetic similarity, especially between E. dubius setosus and E. cil- iatus. One shared feature of possible importance is the elongate, oval, terminal antennal segment. This configuration is considered primitive here, because of its occurrence in related tribes. Within Eusattus it is not a common feature, and could represent a derived condition. The 5 members of the ciliatus subgroup share DOYEN: SYSTEMATICS OF EUSATTUS AND CONISATTUS 19 CO z> _l z> oc III Z) t- < (J CO Z) h- < Q_ O hi CO z> H QQ rr < nr a) ~> Z) _l X (Ji Ql > O Q_ X :: : :: s :: i • (26) ELYTRA, PRONOTAL DISK WOOLY ( 4 ) MENTUM DEEPLY EMARGINATE (55) COXITE LONG (46) HIND TIBIA FIMBRIATE (45) HIND TIBIA BOWED (15) PRONOTUM SQUAMULOSE (27) ELYTRA SQUAMULOSE ( 21 ) ELYTRA NONTUBERCULATE (48) I NTERCOXAL PROCESS BROAD (36) HIND FEMUR SETOSE ( 7 ) ANTENNA SHORT (39) INNER FORETIBIA SPINED (40) INNER FORETIBIA FIMBRIATE (16) PRONOTAL CARINA SETOSE Figure 19. Cladogram of possible relationships among species in the E. muricatus species group. Numbers explained in Appendix A; conventions as in Fig. 13. 3 synapomorphies. Of these, only fimbriate for- etibiae are obviously an adaptation to fossorial habits. The other two apomorphies do not occur elsewhere in Eusattus. The ciliatus subgroup may be divided into two clusters, each distinguished by a pair of apo- morphies: The contiguous foretibial spines shared by E. arenarius, ciliatus, and ciliatoides also oc- cur in the robustus and difficilis species groups. Deep lateral epistomal emarginations (char. 2) do not occur elsewhere in Eusattus, except as aberrations in a few species. Both apomorphies shared by E. pallidus and E. vizcainensis are unique in Eusattus. Distributional Patterns of Eusattus The geographic distributions of the Eusattus species groups in clusters A and B (Fig. 14) are illustrated in Figs. 21 and 22. Distributions of individual species are given in the taxonomic accounts. The largely allopatric distribution of the two major clusters is striking. Cluster A (Fig. 21) is distributed primarily in the semiarid interior re- gions of the southwestern United States and Mexico. The reticulatus species group also occurs on the Pacific slope of central Mexico and in Baja California, with one species nearly reaching the boundary with California. The arid coastal por- 20 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 CO CO CO CO LU z Z> CO Q CO UJ or Z> o Z> 2 CO < -z. < I- < Zj < c_> M CD Z) LU or _l _I _l < Q < O O Q_ > (37) ANTERIOR TIBIA DENTATE (44) FORETIBIA SPINE ROW SHORT (52) COLOR TAN OR BROWN (55) COXITE VERY LONG (53) METENDOSTERNITE ARMS BUTTRESSED (49) METASTERNAL SUTURE SHORT (47) TARSI SETOSE (36,46) HIND FEMUR a TIBIA FIMBRIATE ( 2 ) LAT. EPIST. EMARGINATIONS DEEP (43) FORETIBIAL SPINES CONTIGUOUS ( I ) EPI STOMAL SUTURE DEEP (40) FORETIBIA FIMBRIATE (10) ANTENNAE CLUBBED Figure 20. Cladogram of possible relationships among species in the E. Appendix A: conventions as in Fig. 13. ciliatus species group. Numbers explained in tions of Sonora and Sinaloa do not support species from cluster A. The distribution of the muhcatus species group (cluster A) includes most of the Great Basin, the Mojave and Colorado deserts, and parts of the basin and range province extending into Arizona and New Mexico. Within this area the distri- bution is extremely patchy, corresponding to the occurrence of aeolian sands, so that at most lo- calities muricatus group species are allopatric with species of other groups. The distribution of cluster B species (Fig. 22) is almost entirely along the Pacific slope. The two species of the difficilis group and E. dubius (ciliatus group) occur in transmontane deserts of southern California and Nevada. The depres- sus group occupies the cismontane area of northwestern Mexico, which lacks species of cluster A. The pattern evident in Figs. 2 1 and 22 prob- ably represents a long-standing vicariance. The cluster A species, with the exception of the mur- icatus group, probably evolved in grassland and woodland habitats, which most species now oc- cupy, and are largely active during the summer rainy season as adults. The cluster B species near- ly all inhabit sandy, extremely well-drained sub- strates, and most are largely restricted to sand dunes. The degree of morphological specializa- tion of most of these species suggests a long pe- riod of dune habitation. The occurrence of members of both clusters A DOYEN: SYSTEM ATICS OF EUSATTUS AND CONISATTUS < ID Ld o O > q: h- z Z> UJ o ^ tr o CO c c o — 'j3 T3 2 c c c o J3 -S "S c 2 c ** m 73 .2 1 2 ix p E E t ,o o '— 0 ix jr _3 — u c o ° 2 8. a oo 2 £ c c o ■- c T3 ~ 2! ix 00 -C 3 S 8 o c O c ci -C """ — 1— <« PQ c _ §"« o r) u BO U O C <_ 8. o ix o CQ c «- 2 3 -S C §■.$? o « - > Q. ,x O "" & & = F .2 b § •coo ix eg IS Q <^j I 5 ix O. ,- ■*= U — .c 22 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 and B in the Great Basin may be secondary. Both E. muricatus and E. dubius, which are the only species in large areas of Nevada and Utah, show little geographic differentiation in that region, possibly reflecting relatively recent range expan- sion. However, along the western edge of its range the muricatus group is represented by several other very distinct species, suggesting a longer period of habitation. It may be recalled here that the muricatus group is phenetically very similar to some members of cluster B, especially the cil- iatus group. The concentration of muricatus group species in the Mojave Desert and southern Great Basin coincides with the general distribution of cluster B, and it is possible that the female re- productive tract of the muricatus group has con- verged to resemble that of the reticulatus and convexus groups. In that case, the eastward ex- pansion of the muricatus group into New Mexico and southern Texas would be secondary. Such biogeographic considerations underscore the problems in interpretation of homology and character polarity and the uncertainty of clado- grams. The most interesting distributional patterns are disjunctions across the Gulf of California. These involve the reticulatus group (cluster A) and the depressus group (cluster B). In the reticulatus group, 5 species occur in mainland Mexico or the southwestern United States, and 1 1 occur on the Baja California pen- insula. E. reticulatus is widespread in the south- western United States (Fig. 86), but most of the other species, especially those in Baja California, have restricted distributions. The greatest con- centration of species is in the southern part of the peninsula, and only E. costatus occurs as far north as California. The reticulatus group comprises 3 main lin- eages: E. laevis; the cluster of species related to E. costatus; and the cluster related to E. reticu- latus. The first 2 lineages occur only in Baja Cal- ifornia and are not relevant here. Five of the species related to reticulatus occur on the main- land and 5 on the peninsula. Of the peninsular species, only E. erosus and catavinus occur out- side the Cape region, and are then restricted to a few oases (Fig. 74). Either dispersal or vicariance theories could be invoked to explain the presence of the pen- insular species. Axelrod (1979 and earlier papers) espouses a gradual development of the desert flora of North America. Although semi-arid con- ditions existed by the Miocene, the present re- gional deserts probably developed only during the Pleistocene. In the intervening time, various savannah and dry woodland vegetation domi- nated. If this scenario is correct, Eusattus (and other woodland inhabitants) could have dis- persed across areas that are now extremely arid. As discussed below, the Gulf of California may have been much smaller or nonexistent at this time, so that dispersal need not have been at such a high latitude as at present. Dispersal is considered unlikely for the follow- ing reasons. First, although the cladistic rela- tionships of the reticulatus group species are often uncertain, the peninsular species are definitely not closely related to E. reticulatus. Rather, the most striking character correspondences are be- tween the peninsular species and the mainland cluster pons-venosus-mexicanus. Second, the distribution of the peninsular species suggests a southern origin. If dispersal had been from the north, relictual populations would be expected at other oases along the eastern escarpment of the peninsular ranges. This pattern is seen in species such as Microschatia championi Horn. A vicariance explanation would require iso- lation of the peninsular species by the tectonic events that splintered Baja California from mainland Mexico. Two processes were impor- tant: northward movement of the peninsula rel- ative to the mainland, and opening of the Gulf of California. The northward displacement of Baja California was first suggested by Wegener (1928), solely on the basis of the fit between the opposing coastlines. Phillips (1964) and Larson et al. (1968) suggested an essentially similar movement, which is supported by the pattern of faults and geomagnetic anomalies between the Cape region and the Tres Marias Islands. Ac- cording to these reconstructions, the Cape region originally fit into the embayment between the Tres Marias and Nayarit. The total northward displacement would then amount to approxi- mately 500-600 km. As northward movement occurred, the pen- insula also became separated from the mainland by a seaway which became the present Gulf of California. The Gulf was widened by ocean-floor spreading as well as faulting (Larson et al. 1 968), resulting in a displacement from the continental mass of about 250 km. The timing of these events is disputed. Hill and Dibblee (1953) and Gastil et al. (1975) believe that movement along the DOYEN: SYSTEMATICS OF EUSATTUS AND CONISATTUS 23 San Andreas fault began in Mesozoic time, and Gastifs group and Rusnak et al. (1964) believe that a Gulf seaway existed by the late Miocene. Larson suggests that displacement from the mainland took place largely during the past 4 million years, and that a very narrow seaway, at most, existed before that time. Regardless of exactly when a major seaway appeared, a barrier to dispersal of relatively ses- sile organisms like Eusattus has probably existed for a considerable time. The peninsular moun- tain ranges cast a rain shadow onto the coastal plain of Sonora and northern Sinaloa, and this effect probably extended farther south in the past. The climate of Baja California is also moderated by the cool Pacific Ocean, an influence which the much warmer Gulf of California does not exert. The present thorn forest and scrub vegetation of coastal Sonora and Sinaloa are not nowinhabited by species of the reticulatus group, and would probably prevent dispersal even if no water bar- rier existed. The present distribution of Eusattus coincides remarkably with the geological events described above. E. venosus occurs in Nayarit and Jalisco, and E. mexicanus from Jalisco to Guerrero. In both of these species (and in pons) the pseudepi- pleuron is cristate, at least anteriorly. Cristate pseudepipleura are found elsewhere only in E. catalinensis and ceralboensis, with occur on the indicated Gulf islands. Other characters (Fig. 1 7) show different distributions, and it is uncertain whether the specialized pseudepipleural struc- ture developed once or several times. Initially catalinensis and ceralboensis might have di- verged from a noncristate peninsular stock when their respective islands were isolated. This di- vergence must have been geologically recent, probably no more than a few hundred thousand years, since most of the islands were connected to the peninsula by land-bridges during the Pleis- tocene (Wilcox 1978). In this case, similarities would be convergent. Alternatively, ceralboensis and catalinensis could represent relicts of a cris- tate species which previously occurred on the peninsula. This possibility is suggested by their high degree of similarity and the occurrence of other quite distinct species on intervening is- lands. A quite different vicariance pattern is suggest- ed by the very strong phenetic resemblance of E. pons to E. catavinus and E. erosus. In early literature, pons was several times referred to as erosus (see taxonomic account). The resem- blance to catavinus is even more striking, espe- cially in individuals of pons in which the pseud- epipleural cristae are weak. The distribution of pons largely coincides with the drainage of the southern tributaries of the Rio Grande (Fig. 87). A faunal connection between Baja California and the Rio Grande drainage would seem improba- ble on first examination. However, this pattern recurs in the fossorial, flightless tenebrionid ge- nus Crypt adius. Most species of Cryptadius occur on maritime sand dunes on the Pacific coast, including the Gulf of California. Cryptadius tri- plehorni Berry (1974) inhabits riverine dunes in the Big Bend region of the Rio Grande. The only apparent explanation for this disjunction is that populations inhabiting riverine sands in Sonora reached the Rio Grande drainage by stream cap- ture. It is unknown whether Cryptadius presently occurs in riverine sands in northwestern Mexico. In any case, such a scenario could explain the similarity of pons and catavinus. A cautionary' note must be repeated, that cladistic relation- ships of the reticulatus group species are subject to reinterpretation, and the relationships dis- cussed here are not certain enough to include in Fig. 17. A different pattern is seen in the depressus species group. In contrast to the species of the reticulatus group, which share a relatively high overall similarity, the two species of the depres- sus group are very different in superficial fea- tures. E. secutus is phenetically similar to E. du- bius\ E. depressus is distinctive, without strong resemblance to any other species. Despite their superficial difference, the apomorphies uniting secutus and depressus (Fig. 14) are convincing, and there seems little doubt that they are sister species. The range of depressus encompasses thorn for- est and dry subtropical woodland habitats from northern Nayarit to central Sonora. In Baja Cal- ifornia, secutus occupies similar habitats in the Cape region. Because of their numerous phenetic differences, these species must have been sepa- rate for a long period of time, probably antedat- ing the formation of the Gulf of California, and certainly corroborate a relatively early separa- tion of the peninsula. In other Tenebrionidae. vicariance patterns linking Baja California and west coastal Mexico are not apparent, in part because of the rudi- mentary state of taxonomic knowledge. Some 24 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 27 Figures 23-29. Diagnostic structures of genera of Coniontini. 23. Foretibia and tarsus of Coelus globosus LeConte. 24. Foretibia of Bronchus obscurus. 25. Ventral aspect of left hypomeron of Conisattus rectus, showing marginal setae. 26. Same. Eusallus minimus, showing submarginal setae. 27. Clubbed antenna of Branchus obscurus. 28. Ventral aspect of cranium of E. mexicanus, showing distinct submentum. 29. Same, E. cienegus, with rudimentary submentum. taxa which may show peninsular-mainland re- lationships include the genera Eleodes and Ar- goporis and the tribes Eurymetopini and Asidini. The Epitragini, strongly developed in the sub- tropical areas of Mexico, are unexpectedly de- pauperate in Baja California. One other apparent vicariant relationship needs to be discussed here. The robustus and rudei groups, restricted to the California Channel Is- lands and the Cape region of Baja California, respectively, show an unusual secondary sexual character, described earlier. If this feature, which occurs in one species of each group, is a valid synapomorphy, it represents an enormous dis- DOYEN: SYSTEM ATICS OF EUSATTUS AND CONISATTL'S 25 junction. Closed-cone pines show vaguely sim- ilar distributions, with relictual populations as far south as Cedros Island (see Axelrod 1967). The tenebrionid Coeloenemis has species in southern California and northern Baja Califor- nia, on Cedros Island, and an undescribed species in the Cape region (Doyen 1973). These taxa are northern elements with relictual southern pop- ulations, whereas Eusattus is basically austral. The southern Channel Islands lepidopteran fau- na shows some austral affinities (Powell 1983), but these are mostly desert species generally dis- tributed in northern to central Baja California or occurring even more widely. Key to the Genera of Coniontini 1. Foretarsus with all segments cylindrical, apically truncate 2 Foretarsus with basal segment bearing long spatulate process extending be- yond 2nd segment (Fig. 23) Coelus Eschscholtz 2( 1 ). Foretibia with outer margin expanded as prominent, lamellate process (Figs. 30, 4 1 ) 3 Foretibia subcylindrical to apex, some- times briefly hooked at apex of outer margin (Fig. 24) 4 3(2). Foretarsus with basal segment subequal in length to 2nd segment (Fig. 3); pro- notal carina bearing fringe of long, slender setae (Fig. 25); hypomeron densely setose throughout Conisattus Casey Foretarsus with basal segment more than 2 x length of 2nd segment (Fig. 62); pronotal carina glabrous or with few short, squamose setae; hypomeron glabrous or nearly so, except for fringe of long, slender setae subtending pro- notal carina (Fig. 26) _ Eusattus LeConte 4(2). Antenna with distinct, 3-segmented club (Fig. 27); elytra bearing clavate setae; submentum absent (Fig. 29) 5 Antenna gradually enlarged to apex (Figs. 33, 34), never clubbed; elytra glabrous or bearing slender setae; submentum present (Fig. 28) Coniontis Eschscholtz 5(4). Pronotal disk setose; protibia expanded apically as brief hook (Fig. 24) .... Branchus LeConte and Anectus Horn Pronotal disk without evident setae; protibiae subcylindrical, gradually en- larged to apex Oxinthas Champion Conisattus Casey Conisattus Casey, 1895:614; 1908:146; Gebien 1938:408; Arnett 1960:678; Hatch 1965:138. Coniontis (in part). Doyen 1972:373. Eusattus (in part), Doyen 1977:1. Small, elongate oval, tentyrioid Tenebrioni- dae. Head at rest amplected into thorax about to posterior margin of eyes; frons and epistomum weakly convex, with shallow depression just be- hind epistomal suture; epistomum shallowly, narrowly emarginate medially; eyes reniform, in- dented about lh by epistomal canthus; antennae slender, extending to pronotal base; submentum about 3 x broader than long, subcrescentic. Pronotum about 2 x broader than long; posterior angles obtuse, scarcely or not produced back- ward; hypomeron sparsely tuberculate, sparsely set with short setae throughout; pronotal carina set with fringe of long, slender setae; prosternal process declivous posteriorly, rounded. Elytra moderately convex, without distinct pseudepi- pleuron; epipleuron reaching elytral margin an- teriorly, narrowing and becoming submarginal posteriorly, especially in females; mesosternum scarcely excavate; metasternal suture about xh. length metasternum; intercoxal process angulate, narrowly rounded at apex; abdominal sternites 1-3 sparsely set with fine, setigerous punctures, becoming coarser on sternites 4 and especially 5. Foretibia with apex of outer margin expanded as lamellate fossorial process (Fig. 30); foretarsus with basal segment subequal to 2nd. Aedeagus with apex slender, attenuate; ovipositor with coxite undivided; paraproct about 4 x length of coxite; spermathecal tubes 4, long, slender, tight- ly coiled, unbranched; spermathecal accessory gland with long, slender duct (Fig. 1); meten- dosternite with arms free, not approaching me- sonotal inflections, not fused with mesocoxal in- flections; mesendosternite with arms slender, not flanged basally, extending about % distance to elytral articulations. Type Species. — rectus Casey, monobasic. Conisattus rectus Casey (Figure 30) Conisattus rectus Casey, 1895:614; 1908:146: Hatch 1965: 139; Boddy 1957:189; Rogers et al. 1978 (biology). 26 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 W ', 1 i 7 ri ' T - ' - • i ' v V V T tyto i.^w ''•;■■ ■■ . ';, • ' i v .';" • '\/ v;w- $<>'?',!•,■■■■■ '"/'! Figure 30. Conisattus rectus Casey. Comontis rectus. Doyen 1972:373. Eusattus rectus, Doyen 1977:1. Conisattus nelsoni Boddy, 1957:188, new synonymy. Male. — Elongate oval, slightly convex, dark brown, setose beetles. Head and pronotum with duplex punctation, larger punctures about 1.5 x eye facet diameter centrally, separated by 1-3 puncture diameters, becoming finer on epistomum; smaller punctures '/3-'/4 that size, separated by 1 diameter or less; epistomum slightly indented at lateral sutures; epistomal suture obscured or obliterated medi- ally; antenna subfiliform basally, apical 3-4 seg- ments enlarged as weak club, terminal segment ovoid, scape and flagellum pubescent; antennal segment length ratios as follows: 1.7:0.8:1.5:1.2: 1 .2: 1 .0: 1 .2: 1 .0:0.9:0.9: 1.1; terminal segment 1 .3- 1.4x longer than broad; mentum subtrapezoi- dal, nearly flat, sparsely set with long setae; an- terior corners acute, rounded; anterior margin evenly, moderately concave; submentum slightly broader than base of mentum; gular sutures con- verging anteriorly, subcontiguous at submen- tum. Pronotum 2.0-2.2 x broader than long, lateral margins arcuate, narrowly beaded; corners ob- tuse, anterior rounded, posterior angulate; disk set with shallow, setigerous punctures 2-3 x eye facet diameter, separated by 1-2 puncture di- ameters; setae about lh length 2nd antennal seg- ment, inclined about 45°; smaller punctures about Vt that size, separated by about 1 diameter; pro- sternum opaque, finely, muricately and setiger- ously punctate; prosternal process margined be- tween coxae. Elytra about 1.5 x longer than broad, lateral margins arcuate, more strongly so posteriorly; disk very finely rugulose, set with strongly muri- DOYEN: SYSTEMATICS OF EUSA 1 1 'US AND CONISATTUS 27 cate, setigerous punctures about 1.5 x eye facet diameter anteriorly, becoming tuberculate pos- teriorly; setae as on pronotum; epipleuron with projecting fringe of long slender setae along dor- sal margin of basal half. Femora set with short, declined setae; protibia with rounded apical process extending to apex of 3rd tarsomere (Fig. 31); outer margin set with stout spinules separated by about 1 spine width basally, contiguous in apical '/6; posterior face with spine field in basal V3-V2, inner margin with row of 6-8 long, stout, and several shorter setae; mesotibia and metatibia moderately densely set with long, sharp spines. Aedeagus as in Fig. 32. Female.— Qualitatively indistinguishable from male, except for genitalia; females are slightly stouter and average about 1 5% larger than males. Measurements. — EL 4.5-6.4 mm; EW 3.0- 4.5 mm; PL 1.2-2.0 mm; PW 2.6-4.0 mm; BD 2.3-3.2 mm. Holotype.— From Oregon, Squally Hill (see "Taxonomic Discussion" below); sex not deter- mined (USMNH). Taxonomic Discussion.— The type series, collected by Schwarz, is labelled "Squally HK, Or." Casey (1908:146) reported the label as ". . . Squally Hill, which seems to be in the neigh- borhood of Astoria." All other specimens of Conisattus I have examined are from the semi- arid, sandy region of south-central Washington, and it seems unlikely that the type specimens are from the coastal region. A "Squally Point Light" is situated on the Columbia River in Wasco Co., Oregon, approximately 3 miles northwest of The Dalles, and there is a sand-spit nearby to which the term "hook" could be applied. This is pre- sumably the locality referred to by Schwarz. Among the type series is a female, which is the largest individual examined, and relatively stout. In other respects the Squally Hill specimens do 32 Figures 31-32. Conisattus rectus. 31. Apex of foretibia. 32. Aedeagus, tegmen (left) and median lobe (right). not differ significantly from C. nelsoni Boddy. In particular, the upper surface of rectus is not gla- brous, as stated by Boddy in Hatch (1965:139). The lateral ciliation of the pronotum and epi- pleura is variable in length and density, and color is dependent on age. For these reasons, C. nelsoni Boddy is placed in synonymy under C. rectus Casey. Distribution and Habitat. — Occurs in the subarid portion of eastern Washington, where it inhabits sandy substrates (Rogers et al. 1978). Rogers et al. (1978: 3.10) include northeastern Oregon in the range of Conisattus, but the only specimens I have seen from Oregon are those from the type series. Additional Material Examined. — Oregon: "Squally HK" (2). Washington: Benton Co.: Hanford Reserve. 640' (2), Rich- land (16), and 10 mi NW Richland (1); Paterson (1); Kittitas Co.: Beverly (3); Vantage (4); Walla Walla Co.: Wallula (2). 28 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Eusattus LeConte Eusattus LeConte. 1851:131; 1862:223; 1866:112; Lacor- daire 1859:220; Horn 1870:292; 1894:423; LeConte and Horn 1883:371; Champion 1884:74; Casey 1908:64; Blais- dell 1923:269; Gebien 1938:402; La Rivers 1949:179; Arnett 1960:677; Papp 1961:116; Triplehorn 1968:379; Doyen 1972:373; 1977:5; Tschinkel and Doyen 1976:335 (biology). Discodemus LeConte, 1866:223; Casey 1908:59; 1924:311. Conipinus LeConte, 1866:223; Casey 1908:162; 1924:311; Gebien 1938:403. Nesostes Casey, 1908:58, 162; Cockerell 1939:284; Gebien 1938:401; Arnett 1960:677; Papp 1961:1 16. Megasattus Casey, 1908:62, 162;Blaisdell 1923:265; Gebien 1938:402; Arnett 1960:677; Papp 1961:116. Eusattodes Casey, 1908:64, 162; Gebien 1938:402; Arnett 1960:677; Papp 1961:116. Eusattus (Conipinus), Leng and Mutchler 1927:35; Arnett 1960:677; Papp 1961:116. Eusattus (Eusattus), Leng and Mutchler 1927:35; Arnett 1960:677. Sphaeriont is Casey, 1908:75, 162; Leng 1920:229; Leng and Mutchler 1927:35; Gebien 1938:404, Arnett 1960:677. Coelosattus Blaisdell, 1927:166; Gebien 1938:408; Arnett 1960:678; La Rivers 1969:6. Eusattus (Sphaeriontis), La Rivers 1948:709; 1949:179. Small to moderate tentyrioid Tenebrionidae with robust, ovoid, or subglobular bodies. Male. — Head at rest amplected into thorax about to anterior margin of eyes; frons and ep- istomum weakly convex; epistomum emarginate medially; eyes weakly reniform, barely indented by epistomal canthus; antennae slender, extend- ing about % distance to pronotal base (short in 33 Figures 33-34. Antennae. 33. Eusattus convexus, showing the subglabrous condition typical of the convexus. reticulatus, and muricatus species groups. 34. E. ciliatus, showing the se- tose condition typical of the ciliatus species group. Antennae of the depressus, difficilis, robustus, and rudci groups bear shorter setae (see Figs. 58 and 61). PSEUDEPIPLEURAL RIDGE _- PSEUDEPIPLEURON-" EPIPLEURON PSEUDEPIPLEURON-- - EPIPLEURON 37 EPIPLEURON- 39 Figures 35-40. Elytral structure, ventral aspect of abdo- men and thorax (right) and transverse section through 1st ab- dominal segment (left). 35-36. Eusattus reticulatus, with epi- pleuron broadened basally; the pseudepipleural margin is cristate. 37-38. E. difficilis, with gradually broadened epipleu- ron and rounded pseudepipleural margin. 39-40. E. robustus, without pseudepipleuron. dilatatus); submentum 3-5 x broader than long, subcrescentic or subtrapezoidal, or rarely absent (cienegus). Pronotum 1.8-4. Ox broader than long, posterior angles acute, weakly to strongly produced backward; hypomeron glabrous or nearly so, polished or finely granulate (me.xica- nus), with submarginal fringe of long, projecting setae (Fig. 26) (reduced in laevis); pronotal carina glabrous or set with few squamose setae; proster- nal process subhorizontal, prominent, apically rounded or broadly angulate and abruptly decli- vous. Elytra weakly to strongly inflated, extend- ing laterally far beyond abdominal sternites, usu- ally forming more or less distinct pseudepipleuron DOYEN: SYSTEM ATICS OF EUSATTUS AND CON ISA 11 IS 29 (Figs. 36, 38); epipleuron reaching lateral elytral margin only at base (exception: robustus). Meso- sternum weakly to moderately concave for re- ception of prosternal process; metasternum sparsely punctate, laterally setose; metasternal suture present or absent; intercoxal process an- gulate to broadly rounded or subtruncate; ab- dominal sternites 1-3 sparsely set with simple or duplex punctures, becoming coarser and closer on sternites 4 and 5. Foretibia with apex of outer margin expanded as lamellate fossorial process (Fig. 41); foretarsus with basal segment at least as long as next 3 combined. Metendosternite with arms very long, connected to mesonotal inflec- tions by very short tendons; sometimes fused with mesocoxal inflections; mesendosternite with arms long, slender to short, flanged at base; ae- deagus fusiform or apically attenuate. Female. — Averaging 5-10% larger, usually slightly more rotund. Ovipositor with coxite un- divided, paraproct 2-4 x length of coxite; sper- mathecal tubes 2-4, long, slender, tightly coiled or short, thick; spermathecal accessory gland with long, slender duct (Figs. 7-12). Type Species.— Eusattus: difficilis LeConte, designated by Casey 1908:56; Discodemus: re- ticulatus Say, monobasic; Conipinus: dubius LeConte, designated by Gebien 1938:24; Ne- sostes: robustus LeConte, original designation by Casey 1908:56; Megasattus: erosus Horn, orig- inal designation by Casey 1908:56; Eusattodes: laevis LeConte, monobasic; Sphaeriontis: mur- icatus LeConte, original designation by Casey 1908:56; Coelosattus: fortineri Blaisdell, mono- basic. Remarks. -Horn (1894:349) included E. sculpt us Champion in his list of Coleoptera from Baja California. I have been unable to locate Horn's material representing sculptus, which is a synonym of convexus LeConte, or to determine the species to which Horn was actually referring. Blaisdell (1943) lists E. erosus and costatus from Isla Cedros, but his specimens cannot be located. The few known specimens of cedrosensis are quite distinct, and it is not apparent which species Blaisdell's material represented. Key to the Species of Eusattus 1. Antennae with distal segments bear- ing a few very short setae, appearing subglabrous (Fig. 33) 2 Antennae pubescent (Fig. 34) 5 DORSAL FRINGE -INNER MARGIN OUTER. MARGIN APICAL^ PROCESS Figure 4 1 . Diagram of foreleg of Eusattus, viewed from behind, illustrating terminology used in keys and descriptions. 2(1). Epipleuron narrowing abruptly just behind humerus (Fig. 36) (reticulatus species group) 9 Epipleuron narrowing gradually be- hind humerus (Figs. 38, 40) 3 3(2). Frons and pronotal disk tuberculate or papillate, at least laterally; inner margin of anterior tibia with comb- like row of erect setae (Figs. 96, 1 00) or long, flying setae (Figs. 88, 93); epipleuron with margins subparal- lel except near humerus (muricatus species group) 28 Frons and pronotal disk punctate: in- ner border of anterior tibia with ir- regular row of short spines (Figs. 64- 66, 127); epipleuron gradually nar- rowed from base to apex (as in Fig. 38) 4 4(3). Anterior tibia with row of contiguous or closely spaced spinules extending to apex of outer margin (Figs. 104, 111, 135) 5 Anterior tibia with irregular row of spinules separated by 1-4 spine widths extending no more than % distance to apex along outer margin (Figs. 64-66) (convexus species group) 25 30 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 44 Figures 42-45. Intercoxal processes (42-43) and prosternal structure (44-45). 42, 44. Eusattus ciliatus. 43, 45. E. depressus. 5(1,4). Intercoxal process narrow, angulate (Fig. 42); presternum before coxae shorter than prosternal process (Fig. 44); median metasternal suture present 6 Intercoxal process broad, truncate, or rounded (Fig. 43); presternum be- fore coxae longer than intercoxal process (Fig. 45); median metaster- nal suture absent {depressus species group) 34 6(5). Antenna with terminal segment rounded, nearly symmetrical (Figs. 46,47) 7 Antenna with terminal segment an- gulate, noticeably asymmetrical (Fig. 48) (difflcilis species group) 32 7(6). Antenna with terminal segment long- er than broad (Figs. 34, 46) 8 Antenna with terminal segment broader than long (Fig. 47) (robustus species group) 33 8(7). Anterior tibia with spines extending along outer margin to apex or near- ly so (Figs. 104, 135); pronotum with lateral margins explanate. horizontal, with edge usually up- turned (ciliatus species group) 36 Anterior tibia with spines extending along outer margin no more than 3/4 distance to apex (Figs. 132, 133); pronotum with lateral margins de- clivous (rudei species group) 35 9(2). Elytra with pseudepipleural margin sharply carinate, at least basally (Fig. 35); pseudepipleural surface concave _ 1 0 Elytra with pseudepipleural margin angulately rounded (Fig. 37) or round; pseudepipleural surface convex or flat 1 5 10(9). Pronotum tumid in lateral silhouette (Fig. 49); elytra coarsely, deeply ru- gose pons Triplehorn Pronotum and elytra evenly convex in lateral silhouette (Fig. 50); elytra punctato-rugulose, rugulose, or costate 1 1 11(10). Elytra each with 6 more or less dis- tinct costae, or ecostate and rugu- lose; submentum a distinct, rect- angular sclerite as wide as mentum 1 2 DOYEN: SYSTEM ATICS OF EUSATTVS AND COMSAT 1 1 S 31 48 Figures 46-48. Antcnnal apices. 46. Eusattus duhius setosus. 47. E. robustus. 48. E. difficilis. Elytra each with 2 complete costae, sometimes with 2-3 additional, in- complete costae; submentum ru- dimentary, if present, as wide as gula cienegus, new species 12(1 1). Elytral disk with reticulate sculptur- ing _ 13 Elytral disk with punctato-rugulose sculpturing 1 4 13(12). Elytra with 2 irregular, longitudinal, slightly raised costulae extending 17(16). on to declivity; pseudepipleural margin narrowly explanate in an- terior '4 becoming angulate pos- teriorly venosus Champion Elytra with 6 irregular, longitudinal costulae extending onto declivity; pseudepipleural margin explanate and slightly upturned throughout its length reticulatus (Say) 14(12). Elytra rugulose or alutaceous; hy- pomeron with wispy fringe of setae not reaching lateral margin catalinensis, new species Elytra coarsely punctato-rugose; hy- pomeron with moderately dense fringe of setae extending to lateral pronotal margin ceralboensis, new species 15(9). Hypomeron bearing marginal fringe of setae which extend beyond pro- notal rim; profemur and mesofe- mur bearing dorsal fringes of long, erect setae 1 8 Hypomeron glabrous or with few, short setae which do not exceed pronotal margin; profemur and mesofemur with short, decumbent setae 1 6 Figures 49 16(15). Mentum about 2 x as broad as long; E. reticulatus anterior tibia with distal process extending well beyond apex of bas- al tarsomere, usually to apex of 2nd or 3rd tarsomere; elytral disk retic- ulate mexicanus Champion Mentum about 1 .5 x as broad as long; anterior tibia with distal process extending no further than apex of basal tarsomere; elytral disk ru- gose, rugulose, or smooth 1 7 Pronotal and elytral disks glabrate, with exceedingly fine punctures, barely visible at 50 x laevis LeConte Pronotal disk set with punctures at least as large as eye facets; elytral disk rugose, rugulose, or costate _ 1 9 -50. Lateral silhouettes. 49. Eusattus pons. 50. 32 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 VJ 51 ~ 52 v 53 Figures 51-53. Male genitalia, tegmen (left) and median lobe (right). 51. Eusattus aridus. 52. E. costatus. 53. E. depressus. 18(15). Elytra roughly sculptured — rugose, scabrous, or costate 1 9 Elytra smooth, glabrate, or finely alu- taceous planulus, new species 19(17,18). Elytra bearing very coarse, deep, rounded excavations, especially in lateral areas 20 Elytra finely rugulose, scabrous, or scabrous-costate 2 1 20(19). Body form ovate (0.73 < EW/EL < 0.81); Baja California Sur erosus H orn Body form short ovate (0.79 < EW7 EL < 0.88); Baja California Norte catavinus, new species 21(19). Epipleuron narrowing very abruptly just behind humerus (Fig. 36) 22 Epipleuron narrowing less abruptly behind humerus (as in Fig. 38) araneosus Blaisdell 22(21). Elytra scabrous-costate; pronotum usually widest just behind middle, distinctly wider than elytra at base; aedeagus with apex sharply atten- uate (Fig. 51) aridus, new species Elytra rugose or scabrous, sometimes with faint costulae; pronotum usu- ally widest just before posterior an- gles, subequal in width to elytra; ae- deagus with apex bluntly rounded (Fig. 52) 23 23(22). Cranium with punctures subequal to those of pronotal disk .24 Cranium with punctures distinctly larger, deeper than those on prono- tum _ franciscanus, new species 24(23). Elytra with strong, rounded costae, only faintly rugose; vertex and pro- notal disk simply punctate cedrosensis, new species Elytra with rugae superimposed over costae, which are often obsolete; vertex and pronotal disk with du- plex punctation costatus Horn 25(4). Presternum glabrous or bearing short, decumbent setae (long in individ- uals of convexus from White Sands, N.M.); anterior femora with scat- tered, decumbent setae dorsally (Fig. 136) 26 Presternum bearing long, erect setae; anterior femora bearing dorsal fringe of long, flying setae (Fig. 65) minimus, new species 26(25). Protibia with apical process extending Vi-2/} length of basal tarsomere (Fig. 64); outer margin of protibia entire 27 Protibia with apical process extending beyond basal tarsomere (Fig. 66), usually to apex of 2nd tarsomere; outer margin of protibia dentate obliteratus Champion 27(26). Elytra rugulose, usually with 3-6 un- dulating, irregular, rounded eleva- tions running longitudinally; length (elytra and pronotum) = 7.1-9.5 mm convexus LeConte Elytra punctate or glabrate, occasion- ally with very faint, longitudinal el- evations; length = 8.2-12.2 mm nitidipennis LeConte DOYEN: SYSTEMATICS OF El S I III S AND ( VMS I II I S 33 Figlires 54-57. Labial configurations. 54. Eusattus muricatus. 55. E. phreaiophilus. 56. E. dubius. 57. E. ciliatus. 28(3). Middle and hind tibiae nearly straight, subcylindrical in cross-section; an- tennal scape longer than foretarsus, pedicel about as long as next 2 seg- ments combined 29 32(6). Middle and hind tibia strongly bowed (Figs. 89, 90); hind tibia strongly flattened; antennal scape shorter than foretarsus, pedicel as long as next 3 segments combined dilatatus LeConte 29(28). Elytra tuberculate throughout; frons and vertex without setae 30 33(7). Elytra punctate basally, becoming tu- berculate on declivity; frons and vertex sparsely set with fine, short setae puberulus LeConte 30(29). Elytra set with long, slender, erect se- tae arising from tubercles; prono- tum densely setose in lateral quar- 34(5). ters hirsutus, new species Elytra glabrous or with few, scattered, very fine setae; pronotum glabrous or with few setae near lateral mar- gins 3 1 31(30). Mentum moderately and evenly emarginate (Fig. 54); foretibia with inner fringe of long, slender setae (Fig. 93) (exception: specimens from northern Baja California) muricatus LeConte Mentum deeply, narrowly emarginate (Fig. 55); foretibia with posterior row of short spines (Fig. 100 phreatophilus, new species Elytra evenly convex, never costulate; punctures on frons and pronotum subequal in diameter to eye facets productus LeConte Elytra costulate, sometimes faintly so; punctures on frons and pronotum 2-3 x eye facet diameter difficilis LeConte Elytra strongly reflexed, with epipleu- ron medial to lateral elytral margin except basally (Fig. 38) politus Horn Elytra with only epipleuron reflexed, with epipleural ridge lateral (Fig. 40) robustus LeConte Anterior femur bearing fringe of long, projecting setae dorsally (Fig. 130); anterior tibia broadly explanate, with outer margin convex and pos- terior face granulose, without spi- nation; elytra and pronotum pol- ished, shining; hypomeron setose over entire surface secutus Champion Anterior femur bearing short, ap- pressed setae (as in Fig. 136); an- terior tibia narrowly explanate, with 34 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 Figures 58-61. Epistomal configurations. 58. Eusattus rudei. 59. E. muricatus. 60. E. cilialus. 61. E. depressus. outer margin sinuate and posterior face sparsely spinose (Fig. 1 27); ely- tra and pronotum dull; hypomeron subglabrous except for few short se- tae near lateral margin 37(36). depressus Champion 35(8). Epistomal emargination deep (as in Fig. 59); elytral declivity papillose; protibia with apical process extend- ing to apex of 2nd or 3rd tarsomere crypticus, new species Epistomal emargination shallow (Fig. 58); elytral declivity finely, sparsely punctate and alutaceous; protibia with apical process extending no further than apex of 1st tarsomere rudei, new species 38(37). 36(8). Anterior tibia with external margin armed with row of contiguous or subcontiguous spines, always ex- tending to apex (Figs. 104, 111); mentum with anterior border straight or weakly indented in mid- dle (Fig. 57) 37 Anterior tibia with external margin armed with row of isolated spines, usually ending before apex (Figs. 122, 125); mentum with anterior border emarginate (Fig. 56) 39 Posterior femur bearing dorsal fringe of long, slender setae (Fig. 113); middle and posterior tibiae bearing long, slender setae on mesal surface; metasternal suture no more than xh length of metasternum 38 Posterior femur subglabrous or with few short setae (Fig. 106); middle and posterior tibiae without long setae; metasternal suture extending entire length of metasternum arenarius, new species Basal tarsomere of middle and hind legs set with inclined setae about as long as 2nd tarsomere (Figs. 1 12- 113); PL/PW > 0.43 ciliatus Horn Basal tarsomere of middle and hind legs subglabrous or set with few se- tae about lh length of 2nd tarso- mere; PL/PW < 0.44 ciliatoides, new species DOYEN: SYSTEMATICS OF EUSATTUS AND CONISA II I S 35 39(36). Body subglobose; anterior tibia with posterior margin armed with fringe of erect spines and/or long flying setae (Figs. 122-123, 125) 40 Body ovoid; anterior tibia with pos- terior margin armed with few short, irregular spines or setae (Figs. 1 1 8- 1 1 9) dubius LeConte 40(39). Anterior tibia with row of spines on outer margin extending V2-% dis- tance to apex (Fig. 1 22); dorsum tan or brown pal/idus, new species Anterior tibia with row of spines ex- tending to apex of outer margin or nearly so (Fig. 125); dorsum black vizcainensis, new species Eusattus convexus Species Group Head with simple or duplex punctation; epi- sternum shallowly to moderately emarginate me- dially, scarcely emarginate at lateral epistomal sutures; eye ovoid to weakly reniform; antenna submoniliform, gradually broadened to 10th seg- ment, 1 1th slightly narrower; subglabrous except for sparse, short, decumbent setae; mentum 1 .4- 1.6 x broader than long, anterior corners acute or nearly right-angled, rounded; submentum slightly narrower to slightly wider than base of mentum; gular sutures subparallel or convergent; separated by Va-3/a mentum width anteriorly. Pronotum broadest at base, lateral margins evenly convergent or straighter near posterior angles; anterior angles rounded, about 90°; pos- terior angles acute; disk finely, sparsely punctate, usually more coarsely so anteriorly and laterally; hypomeron polished, longitudinally furrowed, especially over coxae, lateral setal fringe variable; presternum before coxae much shorter than prosternal process, subglabrous to setose; pro- sternal process about 1.4-1.5 x broader behind than between coxae; sparsely to moderately densely and finely punctate. Elytra broadest near or slightly behind middle, evenly arcuate in anterior % , then more abruptly so to apex; pseudepipleural margin broadly rounded, undefined; epipleuron gradually broad- ened from apex to humerus, alutaceous or shal- lowly rugulose, setose in anterior '/2— %; meso- sternum between coxae slightly narrower than prosternal process, shallowly excavate; meta- sternum finely, sparsely punctate, sparsely setose laterally; metasternal suture variable in length; intercoxal process subtriangular, acutely round- ed. Protibia with irregular row of spinules ending before apex of outer margin, separated by 1 to several spine widths; meso- and metatibiae set with sharp, short spines. Metendosternite not fused with metacoxal inflections, fused with metanotum; mesendo- sternite with arms long (convexus, nitidipennis) or short (obliteratus, minimus); spermathecal tubes short, thick (as in Fig. 7); aedeagus with apex bluntly rounded; coxite length: paraproct length = 0.33-0.42. Eusattus convexus LeConte (Figure 62) Eusattus convexus LeConte, 1851:132; 1 866: 1 1 2; Cockerell and Fall 1907:203 (distribution); Kumar et al. 1976:28 (biology); Doyen 1977:5 (synonymy); Lavigne 1980:41 (bi- ology). Eusattus diffialis, Horn 1870:294. Eusattus sculptus Champion, 1892:510; Horn 1894:349.423; Pallister 1954:44, new synonymy. Eusattus congener Casey, 1 908:7 1 . Eusattus acutangulus Casey, 1908:72. Eusattus rotundus Casey, 1908:72. Eusattus subnitens Casey, 1908:72, new synonymy. Eusattus turgidus Casey, 1908:73. Eusattus peropacus Casey, 1908:73 Eusattus acut us Casey, 1908:74. Eusattus quadratus Casey, 1924:31 1. Eusattus subvelutinus Casey, 1924:312. Eusattus woodgatei Casey, 1924:312. Eusattus (Eusattus) woodgatei, Leng and Mutchler 1927:35. Eusattus (Eusattus) subvelutinus, Leng and Mutchler 1927: 35. Convex, broadly oval, black beetles with fur- rowed, shallowly rugose, or shallowly, coarsely punctate elytra. Male. — Frons set with punctures about '/3-1 eye facet in diameter, separated by about 2-4 puncture diameters, usually becoming slightly coarser, denser near epistomal suture and on ep- istomum, sometimes obscure on epistomum; ep- istomum shallowly emarginate, medially; epi- stomal suture narrow, weakly incised; eye reniform, constricted xli-xk by epistomal canthus. Antenna extending to base of elytra; segment length ratios as follows: 3.8:1.6:2.7:1.9:2.1:2.1: 2.2:1.9:1.7:1.4:1.9; terminal segment 1-1.2 x longer than broad, apex acutely angulate or nar- rowly rounded. Mentum with deep submarginal grooves along posterior xh-lU of lateral borders; anterior border shallowly to moderately and ar- cuately emarginate; submentum slightly broader 36 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 14] Figure 62. Eusattus convexus LeConte. than base of mentum; gular sutures nearly par- allel. Pronotum about 2.2-2.3 x broader than long; disk set with punctures about V4-I eye facet in diameter, separated by about 2-6 puncture diameters; hypomeron glabrous except for submarginal fringe of setae extending slightly be- yond lateral border; presternum punctatorugu- lose, sparsely clothed by decumbent setae about as long as 2nd antennal segment. Elytral disk with 4-6 irregular, low, rounded longitudinal ridges, sometimes completely ob- solete except on declivity; ridges impunctate; depressions irregularly set anteriorly with very coarse, shallow punctures bearing minute, de- cumbent setae, often obscured or lost; punctures gradually coalescing posteriorly, producing erod- ed, anastomosing, or reticulate pattern; punc- tures becoming gradually finer and surface ru- gulose or alutaceous laterally; epipleuron 2.5- 4.0 x broader at base than at 3rd abdominal seg- ment, alutaceous, sparsely set with short to long setae in basal half. Profemur and mesofemur set with short, de- cumbent, to moderately long, retrorse setae; pro- tibia with attenuate apical process extending V2- % length of basal tarsomere; outer margin with row of spinules extending Vi-^A distance to apex, separated by 1-2 spinule widths; posterior face with spine field occupying basal '/2-3/4 (Fig. 64a, b); tarsomere length ratios as follows: 5.5: 1.5: 1.3: 1.3:4.2 (pro-); 7.0:3.0:2.3:1.8:5.0 (meso-); 9.0:3.5: 2.6:5.5 (meta-). Female. — Elytra often broadest behind mid- dle, otherwise differs from male as stated in ge- neric description. Measurements. — EL 5.9-10.2 mm; EW 5.2- 8.4 mm; PL 2.0-3.3 mm; PW 4.6-8.0 mm; BD 3.6-6.0 mm. Lectotype Male (MCZ) from vicinity of Long's Peak, Missouri Territory (now Boulder Co., Colo.); 2 syntypes, New Mexico; 2 syntypes, no data. Lectotype Female for sculptus hereby designated from a series of 9 syntypes (BMNH) from Paso del Norte, Chihuahua, Mexico (Hoege). Type Localities.— E. sculptus, Paso del Norte, Chihuahua, Mexico; congener, Las Vegas, N.M.; acutangulus, Colorado; rotundus, north of Fort Collins, Colo.; subnitens, Arizona; turgidus, Kansas; peropacus, Fort Dodge, Kans.; acutus, Logan Co., Kans.; subvelutinus Roswell, N.M. Diagnosis.— Eusattus convexus is similar to E. nitidipennis LeConte, differing as described in the diagnosis for that species. E. convexus is also similar to E. difficilis LeConte, but has punc- tate or eroded elytra (tuberculate in difficilis) and has the row of spinules on the outer protibial margin separate and extending Vi-^A to the apex (nearly contiguous, reaching apex in difficilis). Variation.—/:, convexus shows considerable morphological variation, both within popula- tions and among localities. Elytral sculpturing varies significantly in all substantial collections from a single locality, as does size. This individ- ual variation is superimposed on broad geo- graphic patterns in several features. The largest beetles (7.2 mm < EL < 10.2 mm) occur in southern New Mexico and west Texas, where elytral furrowing is especially evident. The femoral setae tend to be longer in this region than at other localities. These populations correspond to sculptus Champion and subvelutinus Casey. To the north, west, and east, size gradually de- creases, reaching a minimum in north-central New Mexico and southern Colorado (5.9 mm < DOYEN: SYSTEM ATICS OF EL'SATTUS AND CONISATTUS 37 Figure 63. Known distribution of Eusattus convexus. Single records from Missouri. Tennessee, and Orange Co.. Calif., are not plotted. EL < 8.5 mm). Populations from this area cor- Co., N.M., and El Paso Co., Colo., tend to have respond to convexus LeConte and to most of the the elytral sculpturing greatly reduced; they cor- species described by Casey. Individuals from Taos respond to rotundus Casey. South into Chihua- 38 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figures 64-66. Posterior aspect of forelegs. 64a,b. Eusattus convexus, showing variation in spination of posterior face of tibia. 65. E. minimus. 66. E. obliteratus. hua and Durango, Mexico, body size decreases slightly from the maximum in the border region, and the strong elytral furrows usually contain light-colored accretion, producing a distinctive reticulate appearance. Habitat and Distribution (Fig. 63). — Semi- arid or seasonally arid woodland or grassland from about 1000-2500 m elevation, ranging from Cache Co., Utah and central Wyoming south through New Mexico and Texas to Guadalupe de Victoria, Durango, Mexico; west from New Mexico to the Hualupai Mountains, Mojave Co., Ariz., and east to southwestern Missouri and Dallas, Texas. Most specimens are from New Mexico, Colorado, and Arizona, and E. convexus appears to be rare in the peripheral portions of its range. Eusattus minimus, new species (Figure 67) Strongly convex, broadly oval black beetles with coarsely punctate, shallowly rugulose, or eroded elytra. Male. — Head and pronotum set with duplex punctation, small punctures barely visible at 50 x , separated by 2-3 puncture diameters; frons with large punctures about I-IV2 eye facets in diam- eter, separated by 2-3 puncture diameters, be- coming denser along epistomal suture, then shal- low, obscured on epistomum and genae; epistomum weakly to moderately and evenly emarginate; epistomal suture fine, faintly im- pressed; eye elongate oval, anterior margin en- tire. Antenna extending about halfway to elytral DOYEN: SYSTEMATICS OF KL SAULS AND ( OS1S. II It S 39 Figure 67. Eusattus minimus Doyen. base; segment length ratios as follows: 1.5:0.7: 0.8:0.6:0.6:0.6:0.6:0.6:0.6:0.5:0.7; apical seg- ment subrhomboidal, about 1.3 x broader than long with obtusely rounded apex; antepenulti- mate segment about 1.5 x broader than long. Mentum about 1 .6 x broader than long, anterior angles acutely angulate or narrowly rounded, an- terior margin bisinuate, weakly to moderately emarginate in medial half; submentum slightly narrower than base of mentum; gular sutures convergent anteriorly. Pronotum 2.3-2.9 x broader than long; disk with narrow, impunctate median lines, otherwise set with punctures 1-2 eye facets in diameter, separated by about 2-3 puncture diameters cen- trally, becoming slightly coarser, denser, and muricate laterally and anteriorly; in lateral V6, punctures giving rise to inclined setae about as long as 3rd antennal segment; hypomeron with sparse submarginal fringe of long, projecting se- tae along anterior and lateral margins; prester- num moderately densely set with muricate se- tigerous punctures, setae about as long as basal protarsomere, becoming sparser on prosternal process; prosternal process faintly margined or not, about 1 .4 x wider behind than between cox- ae. Elytra widest near middle; disk shallowly ru- gulose or eroded, set with setigerous punctures 2-3 eye facets in diameter, separated by about 2-4 puncture diameters and often obsolescent centrally, becoming smaller, denser, and muri- cate laterally; setae inclined, about as long as 2nd antennal segment centrally, becoming 2-3 x that length near epipleuron; epipleuron about 2-2.5 x broader at base than at 3rd abdominal sternite, rugulose, sparsely set with long, projecting setae in anterior %. Femora with dorsal fringes of long, retrorse, projecting setae; protibia with rounded apical process extending to apex of 2nd or 3rd tarso- mere (Fig. 65); outer margin with row of 5-8 spinules extending about % distance to apex, sep- arated by about 2-3 spinule widths; tibial margin 40 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 entire or serratulate; posterior face with spinules confined to basal V5; tarsomere length ratios as follows: 1.7:0.4:0.4:0.5:1.5 (pro-); 2.3:0.7:0.6:0.6: 1.5 (meso-); 3.0:0.9:0.8:1.7 (meta-). Female. — Differs as stated in generic descrip- tion. Measurements. — EL 4.0-4.8 mm; EW 3.1- 3.8 mm; PL 1.2-1.4 mm; PW 2.8-3.5 mm; BD 2.3-2.9 mm. Holotype Male (CAS) and 62 paratypes (EME) from Nuevo Leon, Mexico: 7 mi E Estan- cia San Roberto, 1 800 m, V-27- 1 98 1 (J. T. Doy- en and J. K. Liebherr); 10 paratypes (EME), 8 mi E Estanica San Roberto, VIII- 1 1-1977 (E. I. Schlinger). Diagnosis.— Eusattus minimus is most simi- lar to E. obliteratus Champion. In minimus, the hypomeron is fringed anteriorly with long, pro- jecting setae; the presternum is set with long, projecting setae; the antennae reach only about halfway to the elytral base; and the apical seg- ment is bluntly rounded. In obliteratus, the hy- pomeron lacks the anterior fringe; the prester- num is set with short, appressed setae; and the antennae reach almost to the elytral base, with the apex acute. E. minimus is superficially sim- ilar to E. puberulus LeConte, differing in having the inner tibial margin armed with 4-5 short spines (row of long, slender setae in puberulus), as well as many other characters. Eusattus nitidipennis LeConte Eusattus nitidipennis LeConte, 1851:133; 1866:112; Horn 1870:294; Champion 1884:75, plate 3; Doyen 1977:6 (syn- onymy). Eusattus brevis Champion, 1884:75; 1892:509. Convex, broadly oval, black beetles with punc- tate elytra. Male. — Frons set with punctures about 1 lh eye facets in diameter, separated by about 2-4 punc- ture diameters, usually slightly coarser and dens- er on epistomum or about epistomal suture; ep- istomum shallowly emarginate; epistomal suture faint or obsolescent; eye reniform, constricted about 'A by epistomal canthus. Antenna extend- ing Vi distance to elytral base; segment length ratios as follows: 2.5: 1 .3: 1 .7: 1 .4: 1 .6: 1 .6: 1 .5: 1 .4: 1.4:1.3:1.6; terminal segment subrhomboidal, about as wide as long, with apex nearly right- angled, narrowly rounded. Mentum about 1.4x broader than long, with submarginal grooves along posterior half of lateral borders, sometimes present as foveae near posterior corners; anterior corners rounded, acute; anterior border moder- ately and angulately emarginate; submentum slightly wider than base of mentum; gular sutures nearly parallel. Pronotum about 2.2 x broader than long, broadest at base; disk set with punctures about 1-1 Vi eye facets in diameter, separated by about 2-4 puncture diameters medially, becoming coarser, denser anteriorly and laterally; hypo- meron glabrous except for submarginal fringe of setae which extend just beyond lateral border; presternum punctato-rugulose laterally, becom- ing sparsely punctate medially, sparsely clothed with short, decumbent setae; prosternal process unmargined, finely, sparsely punctate, about 1 .4 x broader behind than between coxae. Elytral disk alutaceous or finely rugulose, set with punctures about 1 V2— 2 eye facets in diam- eter, separated by about 2-8 puncture diameters, becoming larger, shallower, or obsolescent on de- clivity; 2-3 very faint longitudinal elevations oc- casionally visible; epipleuron about 2.4-2.9 x broader at base than at 3rd sternite, glabrous or sparsely set with moderately long setae basally. Profemur and mesofemur set with short, de- cumbent setae or occasionally with moderately long, retrorse setae; protibia similar to that of convexus (Fig. 64a), with attenuate apical process extending Vi-V* length of basal tarsomere; outer margin with row of spinules extending %-% dis- tance to apex, separated by 1-2 spinule widths; posterior face with spine field occupying ap- proximately basal half; tarsomere length ratios as follows: 3.6: 1.1:1.1:1 .0:3.5 (pro-); 5.5:2. 1:1.8: 1.5:3.8 (meso-); 7.2:2.7:2.0:3.9 (meta-). Female. — Differs as stated in generic descrip- tion. Measurements. — EL 6.1-9.3 mm; EW 4.8- 7.5 mm; PL 2.1-2.9 mm; PW 4.7-6.5 mm; BD 3.6-5.1 mm. Holotype Male(?) in MCZ. Lectotype Fe- male for brevis hereby designated from a series of 1 3 syntypes (BMNH) from Esperanza, Mexico (Hoege). Additional syntypes in AMNH (4) and USNM(l). Type Localities. — E. nitidipennis, Jalapa, Veracruz, Mexico; E. brevis, Esperanza, Puebla, Mexico. Diagnosis. — Eusattus nitidipennis is most similar to convexus LeConte, but is relatively more elongate (0.75 < EW/EL < 0.80 in niti- DOYEN: SYSTEM ATICS OF El V till S AND ( ONIS. I TTUS 41 MINIMUS OBLITERATUS NITIDIPENNIS Figure 68. Known distribution of Eusattus minimus, obliteratus, and mtidipennis. dipennis\0.8Q < EW/EL < 0.93 in convexus). In nitidipennis the elytra are finely punctate, with 2-3 extremely faint raised lines visible in one individual. In convexus the elytra show 5 or 6 distinct, rounded, longitudinal ridges (reduced or absent in individuals from northern New Mex- ico, Colorado, Utah, and Wyoming). In nitidi- pennis the basal and terminal segments of the protarsus are subequal in length; in convexus the basal segment is about 1 .3-1.5 x longer than the terminal. E. nitidipennis is also similar to E. ob- literatus Champion, differing as indicated in the diagnosis of that species (below). Additional Material Examined (Fig. 68). — Mexico: Coa- huila: South of Saltillo (Rt. 57), VN-10-1963 (1); 16 km SE Saltillo (Rt. 57). V-27-1981 (1). Nuevo Leon: 1 mi E Estancia San Roberto, V-27- 1981(1). Zacatecas: 3 mi W Zacatecas (2); Zacatecas (1). Guanajuato: Guanajuato (2). Veracruz: Espe- ranza (15); Las Vigas (2). Eusattus obliteratus Champion Eusattus obliteratus Champion, 1892:510. Eusattus obsoletus Champion, 1892:51 1, new synonymy. Convex, broadly oval, black beetles with faint- ly furrowed elytra. Male. — Frons opaque, set with shallow, ob- scure, or nearly obsolete punctures about as large as eye facets, separated by about 1-3 puncture diameters, becoming slightly denser about epi- stomal suture; epistomum moderately, narrowly and evenly emarginate (as in Fig. 60); epistomal suture fine, faintly impressed; eye weakly reni- form, barely constricted by epistomal canthus. Antennae extending about 7/x distance to elytral 42 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 base; segment length ratios as follows: 2.0:1.0: 1.4:1.2:1.3:1.4:1.2:1.2:1.2:1.0:1.3; terminal seg- ment about 1 .3 x longer than broad, apex acute. Mentum about 1.6—1.7 x broader than long; an- terior corners broadly rounded, acute; anterior border deeply, evenly emarginate; submentum slightly narrower than base of mentum; gular su- tures convergent anteriorly. Pronotum 2.1-2.5x (usually 2.1-2.3x) broader than long, broadest at base; disk set with weakly muricate punctures about as large as eye facets, separated by about 2-3 puncture diame- ters centrally, becoming denser anteriorly and laterally; hypomeron glabrous except for sub- marginal fringe of setae extending well beyond lateral border; presternum sparsely, very finely punctate, clothed with short appressed setae; prosternal process faintly margined, occasionally unmargined, with a few fine punctures; about 1.5 x broader behind than between coxae. Elytral disk shallowly rugulose, set with punc- tures about 1 Vi eye facets in diameter, separated by about 2-8 puncture diameters and often ob- solescent centrally and on declivity, becoming denser, larger, distinct, and muricate laterally; 5- 6 faint, broadly rounded longitudinal elevations on each elytron, sometimes obsolete; epipleuron about 3-4 x broader at base than at 3rd sternite, alutaceous, sparsely setose in anterior half. Profemur and mesofemur with sparse dorsal fringe of retrorse setae; protibia with attenuate apical process extending approximately to apex of 2nd tarsomere (as in Fig. 66); outer margin with row of spinules extending about Vi-Va dis- tance to apex; spinules separated by about 2-3 spinule widths and subtended by denticulate pro- cesses; posterior face with spinules occupying ap- proximately basal Vi, or extending about halfway along margins; tarsomere length ratios as follows: 2.0:0.5:0.5:0.5:1.8 (pro-); 2.9:1.1:0.9:0.7:2.1 (meso-); 4.3:1.4:1.0:2.4 (meta-). Female. — Differs as stated in generic descrip- tion. Measurements. — EL 5.3-7.1 mm; EW 4.0- 5.6 mm; PL 1.8-2.4 mm; PW 3.8-5.5 mm; BD 3.0-4.2 mm. Lectotype Female for obliteratus hereby des- ignated from a series of 13 syntypes (BMNH) from Villa Lerdo, Durango, Mexico (Hoege). Diagnosis. — Eusattus obliteratus is similar to E. nitidipennis LeConte. In obliteratus, the apical protibial process extends at least to the apex of the basal tarsomere and the outer tibial margin is dentate (Fig. 66); the hypomeral setae are long- er, extending well beyond the pronotal margin. In nitidipennis, the protibial process extends about Vi-Va the length of the basal tarsomere, the outer tibial margin is entire, and the hypomeral setae extend barely beyond the pronotal margin. E. obliteratus is also similar to E. minimus Doy- en, differing as indicated in the diagnosis of that species. Additional Material Examined (Fig. 68). — Mexico: Coa- huila: 20.9 km S Cuatro Cienegas, X- 1-1 976, gypsum dunes (7); 28 mi W Paila, VII-15-1975 (17); 52 mi W Saltillo, VII- 15-1975 (1). Eusattus reticulatus Species Group Moderately to strongly convex, elongate oval to broadly oval beetles with variably sculpted elytra. Head and pronotum with simple or duplex punctation, epistomum shallowly to moderately emarginate medially, scarcely to moderately emarginate at lateral epistomal sutures; antenna submoniliform, gradually broadened to 9th or 10th segment; scape and basal few segments sparsely set with setae V2-I x length 2nd segment, becoming subglabrous distally (entire flagellum sparsely setose in laevis). Pronotum broadest at base (broadest well be- fore base in aridus), lateral borders arcuate or nearly straight in posterior half, then evenly con- vergent to rounded, nearly right-angled anterior corners (exceptions: mexicanus, venosus); pos- terior corners strongly produced posteriorly, acute; hypomeron glabrous except for submar- ginal fringe of setae; presternum before coxae much shorter than prosternal process; prosternal process usually unmargined (variable in erosus), very sparsely, moderately coarsely punctate, or with duplex punctation. Elytral base subequal in width to pronotal base (narrower in aridus), sides arcuate, widest near middle, or nearly straight in anterior half {ari- dus), then convergent to apex; pseudepipleural margin rounded, angulate, or explanate; epipleu- ron as wide as pseudepipleuron at base, imme- diately narrowed to '/z-'/a that width, thence sub- parallel nearly to apex or more gradually narrowed (araneosus); mesosternal width be- tween coxae subequal to width of prosternal pro- cess (exception: laevis); intercoxal process acute, narrowly rounded. Foretibia with irregular row of spinules extending '/2-% distance to apex; mid- dle and posterior tibiae set with short, sharp DOYEN: SYSTEMATICS OF EUSATTUS AND CONIS II II \ 43 spines. Metendosternite not fused with coxal in- flections, fused with metanotum; mesendoster- nite arms extending about % distance to notum; spermathecal tubes several, short, thick (Fig. 7); aedeagus with bluntly rounded apex (sharply at- tenuate in aridus); ovipositor with coxites nearly straight, apex bluntly rounded; coxite length- paraproct length ratio variable. Eusattus araneosus (Blaisdell) Megasattus araneosus Blaisdell, 1923:266; 1943:191. Convex, broadly oval, black beetles with ru- gulose elytra and relatively broad epipleuron. Male. — Head and pronotum with duplex punctation; frons with large punctures about lh eye facet in diameter, separated by 1-4 puncture diameters, becoming denser above eyes and on epistomum; small punctures about Vs that size, separated by about 1 puncture diameter; epi- stomal suture largely obliterated medially by cra- nial sculpturing; epistomum moderately emar- ginate; antennal segment length ratios as follows: 2.6:1.0:1.9:1.3:1.4:1.4:1.3:1.2:1.2:1.0:1.3; ter- minal segment 1.2 x longer than broad, scape and flagellum subglabrous; mentum 1.5 x broad- er than long, subglabrous, anterior corners acute, broadly rounded, anterior border deeply, evenly emarginate, lateral borders weakly reflexed in posterior quarter. Pronotum 2.2-2.4 x broader than long, broad- est at base; disk with larger punctures about as large as eye facets centrally, separated by 2-4 puncture diameters; small punctures V3-V2 that size, separated by 1-3 puncture diameters, lateral margins moderately explanate, weakly rugulose; hypomeron glabrous except for moderately dense submarginal fringe of setae extending well be- yond border; sternum rugulose, sparsely set with short, declined setae; prosternal process finely punctato-rugulose, glabrous. Elytral disk finely, evenly rugose, noncostate; pseudepipleural margin obtusely rounded; pseudepipleuron flat, finely rugose; epipleuron narrowed abruptly behind humerus, equal to width of pseudepipleuron at middle of 1st ster- nite, thence narrowing very gradually, subpar- allel to apex; sparsely set with long, projecting setae in basal half. Anterior 3 abdominal ster- nites muricately punctate, last 2 simply punctate. Profemur and mesofemur with sparse dorsal fringe of moderately long, inclined setae; protibia with apical process extending to apex of 1 st tar- somere; outer margin with row of spinules ex- tending about 3/4 distance to apex, separated by 1-4 spinule widths: tarsomere length ratios as follows: 3.4:0.7:0.6:0.6:2.7 (pro-); 4.5:1.4:1.3:1.2: 3.2 (meso-); 6.3:2.0:1.4:3.5 (meta-). Female. — Differs as stated in generic descrip- tion; ovipositor not examined. Measurements. — EL 10.9-1 1.7 mm; EW 8.3- 9.2 mm; PL 3.4 mm; PW 7.6-8.0 mm; BD 5.8- 6.1 mm. Holotype Female and Allotype Male (CAS) Santa Inez Island, Baja California Sur, Mexico, V- 13- 1921 (E. P. Van Duzee). Diagnosis.— Eusattus araneosus is similar to E. franciscanus Doyen in dorsal sculpturing. It differs from franciscanus and all other members of the reticulatus species group in the shape of its epipleuron, which is relatively broad and not as abruptly narrowed behind the humerus. Eusattus aridus, new species (Figure 69) Convex, broadly oval black beetles with cos- tate, rugose elytra. Male. — Head and pronotum with duplex punctation; frons set with large punctures about 1-2 eye facets in diameter, separated by about 1-3 puncture diameters, becoming coarser and separated by 1 diameter on epistomum; small punctures about V4-V6 that size, separated by about 1 puncture diameter; medial epistomal suture weakly impressed or obscured; epistomum shal- lowly, usually angulately emarginate; terminal segment acutely angulate; antennal segment length ratios as follows: 3.5:1.8:3.6:2.7:2.6:2.5: 2.3:2.1:2.0:1.8:2.4; terminal segment 1. 1-1.4 x (2.4 x in one specimen) longer than broad, acute- ly angulate; mentum about 1.4 x broader than long, almost flat, sparsely setose, anterior corners acute, rounded; anterior border deeply, angu- lately emarginate; submentum as wide as base of mentum; gula anteriorly about lh width of submentum. Pronotum 2.0-2.6 x broader than long, usu- ally broadest just behind middle, with sides evenly arcuate; occasionally broadest near base, with sides nearly straight in middle third; disk set cen- trally with punctures about as large as eye facets, separated by 2-4 puncture diameters, becoming 2 eye facets in diameter and denser laterally; lat- eral margins strongly explanate, punctato-rugu- lose; hypomeron longitudinally furrowed, gla- brous except for moderately dense submarginal 44 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 Figure 69. Eusatlus aridus Doyen. fringe of setae which project beyond lateral mar- gins; sternum punctato-rugulose to scabrous, sparsely set with very short, fine setae; prosternal process 1.5 x wider behind than between coxae. Elytra distinctly narrower than pronotum at base, subparallel in anterior half, then conver- gent; disk costate-rugose or costulate-rugose, usually with 5 distinct shining ridges on each side interconnected by irregular rugae; intercostal depressions opaque, sparsely set with minutely setigerous punctures; pseudepipleural margin obtuse or right-angled, narrowly or broadly rounded; pseudepipleuron flat, punctato-rugose to coarsely rugose; anterior 3 sternites rugulose, sparsely, muricately punctate, or asperate; last 2 sternites coarsely, more densely punctate or punctato-rugulose. Profemur and mesofemur with sparse to mod- erately dense fringes of long, erect or retrorse setae; protibia with apical process extending to apex of 1st or 2nd tarsomere, outer margin with row of spinules extending %-% distance to apex; separated by 1-3 spinule widths (Fig. 78); tar- somere length ratios as follows: 6.4:1.8:1.6:1.5: 5.6 (pro-); 7.0:3.4:2.6:2.4:7.0 (meso-); 11.3:3.7: 2.6:7.0 (meta-). Female. — Pronotal base width subequal to basal elytral width; coxite length ratio to para- proct length = 0.33. Measurements. — EL 10.7-15.7 mm; EW 7.1- 11.6 mm; PL 3.7-4.9 mm, PW 7.8-11.2 mm; BD 5.3-8.3 mm. Holotype Male (CAS) and 28 paratypes from Mexico, Baja California Sur, 1 1.3 km N Guer- rero Negro, IV- 10/1 1-1 976 (R. L. Aalbu; RLA and EME); 3 paratypes, same data, VI-25-1977; 21 paratypes, same data, VII-6-1979. Diagnosis.— Eusattus aridus is similar to E. costatus Horn, but is more strongly costate. In males of aridus the pronotum is distinctly broad- DOYEN: SYSTEM ATICS OF EUSATTUS AND CONISATTUS 45 Figure 70. Known distribution of Eusattus aridus. er than the elytral base (subequal in costatus), and the aedeagus is sharply attenuate (rounded in costatus). Distribution (Fig. 70) and Habitat.— All collections of E. aridus are from the Vizcaino Desert, and all are from sandy substrates, usually dunes. Sympatric populations of costatus and ar- idus are unknown, but the two species occur within about 1 5 miles of one another on the coast north of Guerrero Negro and near San Ignacio on the eastern edge of the Vizcaino. Intermediate individuals are not known. The adults have been collected in pitfalls, from the sand surface at night, and from the sand beneath Encelia bushes. Additional Material Examined. — Mexico: Baja Califor- nia Sur: 2 mi S Ejido Morelos. VII-5-1977 (3); 5 mi SW Jet. Fisher's Cafe, VII-3-1977 (2); 14 mi S Ei Arco. VI- 16- 1967 (1); 5.5 mi SE Millers Landing, VII-6-1977 (2); 6 mi N Guer- rero Negro, VII-4-1979 (31); 7 km N Guerrero Negro, 111-21- 1975 (12); 26.5 mi SE Guerrero Negro, IH-19-1981 (5). Baja California Norte: Rancho Mesquital, VI- 14/ 15- 1967 (1). Eusattus catalinensis, new species Convex, broadly oval, black beetles with finely eroded, carinulate elytra with the pseudepipleu- ral margin carinate. Male. — Head and pronotum with duplex punctation; frons set with large punctures about 1-2 eye facets in diameter, separated by 2-3 puncture diameters; small punctures about lA that size, nearly confluent, especially in epistomal re- gion, producing granulate appearance. Epistomal suture obscured or obliterated medially by cra- nial sculpturing; epistomum weakly to moder- ately, evenly emarginate. Antennal segment length ratios as follows: 4.0:1.9:2.8:2.2:2.2:2.0: 2.0: 1 .8: 1.5: 1.5:2.3; terminal segment about 1.3 x longer than broad, with acutely angulate apex. Mentum about 1.4 x broader than long, granu- late, with few short, appressed setae, anterior cor- ners acute, broadly rounded; anterior border deeply, angulately emarginate; submentum about as wide as base of mentum; gula anteriorly about l/3 width of submentum. Pronotum about 2.3-2.4 x broader than long; disk with large punctures subequal to eye facet in diameter, separated by 2-4 puncture diame- ters medially, becoming denser anteriorly and laterally; small punctures separated by about 1 puncture diameter; hypomeron glabrous or with exceedingly fine, sparse submarginal fringe of se- tae not approaching pronotal border; sternum rugulose-granulate, bearing few short, fine setae and appearing glabrous; prosternal process un- margined, densely, finely punctate or punctato- granulose. Elytral width at base subequal to pronotal width; disk set with numerous irregular, shallow erosions, coarser and somewhat coalescent along lateral margin; each elytron usually with 4-5 faint, smooth carinulae nearly reaching apex; pseud- epipleural margin narrowly explanate, upturned (as in Fig. 35); pseudepipleuron concave, aluta- ceous or finely wrinkled. Metasternum with du- plex punctation, asetose; abdominal sternites with duplex punctation; small punctures nearly con- tiguous on last sternite, which appears granulose. Profemur and mesofemur with short, declined setae dorsally; protibia with narrowly rounded apical process extending almost to apex of basal protarsomere (as in Fig. 78); outer margin with irregular row of spinules extending %-7/8 distance to apex, separated by 1-2 spinule widths; tar- somere length ratios as follows: 5.3:1.1:1.2:1.3: 46 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figure 71. Eusattus catavinus Doyen. 4.5(pro-);7.2:2.1:1.9:1.5:5.6(meso-);9.8:3.2:2.4: 6.0 (meta-). Female.— Coxite length ratio to paraproct length a 0.45. Measurements. — EL 9.4-14.1 mm; EW 7.8- 1 1 .2 mm; PL 2.8-4.2 mm; PW 6.4-9.9 mm; BD 5.2-7.7 mm. Holotype Female (CAS) and 6 paratypes (EME) from Isla Santa Catalina, Baja California Sur. Diagnosis. — Eusattus catalinensis is most similar to E. ceralboensis Doyen, with which it shares the explanate pseudepipleural margins. The latter is much more coarsely sculptured and has the hypomeral fringe of setae much denser and reaching the pronotal border. E. pons Tri- plehorn, E. reticularis (Say), and E. cienegus Doyen have the pseudepipleural margin expla- nate, but differ in numerous other features and occur in the southwestern United States and on the central plateau of Mexico. Discussion.— One specimen collected by J. P. Figg-Hoblyn and labeled Isla Santa Catalina ap- pears conspecific with E. erosus laeviventris (Blaisdell), which is otherwise known from Isla Santa Cruz. Figg-Hoblyn collected on both is- lands on consecutive days, and it seems very likely that the locality data were confused. Additional Material Examined. — Mexico: Baja Califor- nia Sur: Isla Santa Catalina: 111-27-1953 (9 damaged speci- mens); IV-3/4-1974 (9 damaged specimens); W side, VI-25- 1964 (1 damaged specimen). Eusattus catavinus, new species (Figure 71) Strongly convex, very broadly oval black bee- tles with coarsely rugose elytra. Male. — Head with duplex punctation; frons DOYEN: SYSTEMATICS OF EUSATTLS AND ( OMSAIITS 47 set with large punctures about 2-3 eye facets in diameter, separated by about 1-3 puncture di- ameters, becoming denser above eyes and coarser and almost contiguous near epistomal suture; small punctures about '/» that size, separated by about 1 puncture diameter. Epistomal suture faint, often obliterated medially by cranial sculp- turing; epistomum moderately and evenly emar- ginate. Antennal segment length ratios as follows: 3.4:1.8:2.7:1.8:1.8:1.8:1.6:1.7:1.7:1.4:1.6; ter- minal segment about as long as wide, apex an- gulate, mentum about 1.5-1.6 x broader than long, subglabrous; anterior corners acute, broad- ly rounded; anterior border deeply, evenly, or angulately emarginate; submentum slightly broader than base of mentum; gula anteriorly about 0.4 x submentum width. Pronotum 2.2-2.6 x broader than long; disk with duplex punctation; large punctures 1-2 eye facets in diameter, densest laterally and ante- riorly; small punctures about Vt that size, densely, evenly distributed; hypomeron glabrous except for sparse marginal fringe of setae extending just beyond pronotal border; sternum rugulose, sparsely setose; prosternal process margined or not, punctate or rugulose anteriorly, becoming nearly glabrous posteriorly. Elytral width at base subequal to pronotal width; disk coarsely rugose or cariose, especially near lateral margins, noncostate; lateral elytral margin slightly obtuse in cross-section, broadly rounded; pseudepipleuron feebly to moderately concave, strongly rugose. Metasternum moder- ately coarsely punctate medially, becoming mu- ricately punctate and setose laterally; anterior 3 sternites with duplex punctation. Profemur and mesofemur with sparse dorsal fringe of moderately long, erect or retrorse setae; protibia with sharply attenuate apical process ex- tending about to apex of basal tarsomere (as in Fig. 78); outer margin with row of spinules ex- tending about 2h distance to apex, separated by about 1 spinule width; tarsomere length ratios as follows: 4.6: 1.5: 1.3: 1.4:4.6 (pro-); 7.1:2.8:2.5:1.9: 5.2 (meso-); 9.7:3.8:2.8:6.1 (meta-). Female. — Coxite length ratio to paraproct length s 0.45. Measurements. — EL 9.3-12.7 mm; EW 7.7- 10.8 mm; PL 2.9-4.2 mm; PW 7.0-9.4 mm; BD 5.3-7.4 mm. Holotype Female (CAS) and 21 paratypes (EME) from Mexico, Baja California Norte, Ar- royo Catavina, 35 mi SE El Progresso, 2-IV- 1 976, J. D. Lot 76D1. Additional paratypes as follows: 10 mi N Catavina, 4-V-1977, R.'Selb (9, CAS); 5 km N Catavina, 9-IV-1976, R. L. Aalbu (5, RLA); 2.1 mi NW Catavina, 3-V1I-1979, R. L. Aalbu (2, RLA); 1 mi SE Catavina, 3-VII-1977, F. G. Andrews and A. R. Hardy (2, CDFA); 59 mi SE San Quintin, 111-20-1975, R. L. Aalbu (3, RLA). Diagnosis.— Eusattus catavinus is most sim- ilar to E. erosus Horn, which differs in its more slender hind body, sparser, shorter femoral and hypomeral setal fringes, and finer elytral rugosity (see erosus for details). E. catavinus is also sim- ilar to E. pons Triplehorn, but in pons the pseud- epipleural margin is carinate (broadly rounded in catavinus). E. catavinus is superficially similar to some specimens of costatus Horn, differing as stated in the diagnosis for the latter. Distribution (Fig. 74) and Habitat. — Known mainly from the palm oasis at Arroyo Catavina. The beetles inhabit the sandy canyon bottom, where they aggregate in the litter beneath the canopies of shrubs such as Encelia. Additional Material Examined. — 7 damaged specimens with the same data as holotype; 7 with same data as various paratypes; 0.6 mi NW Rancho Santa Ynez. VI- 1 5- 1 975 ( 1 ); 2 mi NW Rancho Santa Ynez, 111-27-1973 (2); Rancho Santa Ynez, no date (1). Eusattus cedrosensis, new species Convex, oval to broadly oval beetles with cos- tate elytra. Male.— Frons set with punctures about lh-2 eye facets in diameter, separated by about 2-4 puncture diameters, slightly denser on episto- mum; epistomum moderately and narrowly emarginate medially; epistomal suture faint, ob- scured medially; mentum about 1.5 x broader than long, slightly convex, anterior corners acute, broadly rounded; anterior border deeply, angu- lately emarginate; lateral borders with submar- ginal groove in posterior fifth; submentum slight- ly wider than base of mentum; gula anteriorly about xh submentum width. Pronotum 2. 1-2.3 x broader than long; disk set centrally with punctures about as large as eye facets, separated by 2-6 puncture diameters, be- coming coarser, denser laterally; presternum ru- gulose; prosternal process coarsely punctate, 1 .5- 1.7 x broader behind than between coxae. Elytra widest near middle, lateral margins ar- 48 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 cuate in anterior half, then more abruptly con- vergent to apex; disk bearing 6 rounded costae plus sutural costa on each side; intercostal fur- rows gently rugose medially, becoming more coarsely so laterally; pseudepipleural margin nearly right-angled, narrowly rounded (as in Fig. 38); pseudepipleuron flat, rugulose; epipleuron rugulose. Metasternum punctato-rugose; ab- dominal sternites 1-3 moderately coarsely punc- tate, becoming rugulose laterally. Aedeagus with bluntly rounded apex. Female. — Differs as stated in generic descrip- tion; ovipositor not examined. Measurements. — EL 9.7 to about 1 1 mm;EW 7.8-10.0 mm; PL 3.3-3.7 mm; PW 6.9-8.5 mm; BD 5.1-6.4 mm. Holotype Male (CAS) from Mexico, Baja California, NE end Cedros Island, 111-27-1952, J. P. Figg-Hoblyn. Paratype (CDFA) from Ce- dros Island, North Point, III-20/2 1 - 1 98 1 , F. An- drews and D. Faulkner. Diagnosis. —Eusattus cedrosensis is similar in dorsal sculpturing to E. aridus Doyen and E. costatus Horn, differing in its more deeply emar- ginate epistomum, the simple punctation of the head and pronotum, and its much more weakly rugose elytra. This species is known only from two partial specimens, largely lacking setae, and the female lacking genitalia. The body shape, epistomal emargination, and abdominal sculpturing are more similar to those features in E. erosus than to the populations of aridus and costatus which occur on the peninsula adjacent to Cedros Island. Blaisdell ( 1 943: 1 90, 191) lists Eusattus erosus and E. costatus from Cedros Island; the speci- mens referred to cannot be located in CAS. Eusattus ceralboensis, new species Megasattus erosus, Blaisdell 1923:265 (in part); 1 943: 190 (in part). Convex, broadly oval black beetles with coarsely eroded, carinulate elytra with the pseud- epipleural margin carinate. Male. — Head and pronotum with distinctly duplex punctation; frons set with large punctures 2-3 eye facets in diameter, becoming denser an- teriorly, sometimes nearly confluent near epi- stomal suture; small punctures about lA that size, evenly separated by about 1 puncture diameter. Epistomal suture faint, usually obscured by cra- nial sculpturing; epistomum moderately to deep- ly, narrowly or angulately emarginate (as in Fig. 60). Antennal segment length ratios as follows: 4.5:2.0:3.4:2.8:2.8:2.8:2.7:2.6:2.4:2.0:2.4; ter- minal segment about as long as broad, with apex nearly right-angled; mentum about 1 .6 x broader than long, sparsely punctate; anterior corners acute, narrowly rounded, anterior margin mod- erately to deeply, evenly emarginate; submen- tum about as wide as base of mentum; gula an- teriorly about Vi width of submentum. Pronotum about 2.3-2.4 x broader than long; disk with large punctures about 1 {h eye facets in diameter, separated by about 2-4 puncture di- ameters medially, becoming coarser, denser an- terolaterally; small punctures no more than about xk that diameter, often barely visible at 50 x, separated by 1-2 puncture diameters; hypome- ron with sparse submarginal fringe of setae ex- tending just beyond pronotal border; sternum punctato-rugulose, sparsely set with moderately long, declined setae; prosternal process with few large punctures anteriorly, finely, densely punc- tate posteriorly. Elytral disk set with numerous deep, irregular excavations, coarser laterally, becoming coales- cent near pseudepipleural margin and usually forming a distinct, punctato-granulate trough; each elytron with 3-4 faint carinulae, scarcely evident on some specimens; pseudepipleural margin narrowly explanate, upturned (Fig. 35); pseudepipleuron concave, irregularly eroded, sparsely set with moderately long setae near base. Metasternum with duplex setation, sparsely set with moderately long setae laterally; abdominal sternites with duplex punctation. Profemur and mesofemur with sparse fringe of short, declined setae dorsally; protibia with narrowly rounded apical process extending to apex of basal tarsomere (as in Fig. 78); outer margin with irregular row of spinules extending about 3/» distance to apex, separated by 1-3 spi- nule widths; tarsomere length ratios as follows; 6.8:1.9:1.5:1.3:5.8 (pro-); 8.6:3.4:2.9:2.6:6.3 (meso-); 11.7:4.3:3.4:6.5 (meta-). Female. — Coxite length ratio to paraproct length s 0.45. Measurements. — EL 1 1.9-14.0 mm; EW 9.4- 12.0 mm; PL 3.6-4.5 mm; PW 8.6-10.3 mm; BD 6.2-7.6 mm. Holotype Female and 5 paratypes (CAS) from Mexico, Baja California Sur, SE end Isla Ceralbo, 111-20- 1 953, J. P. Figg-Hoblyn; 4 paratypes (CAS, DOYEN: SYSTEMATICS OF EUSATTLS AND COXISATTLS 49 Figure 72. Eusattus cienegus Doyen. EME), Isla Ceralbo, Bufo Ranch, 111-22-1953, J. P. Figg-Hoblyn; 2 paratypes (CAS), So. end Cer- ralvo Island, IV- 1-1 952, J. Figg-Hoblyn; 6 para- types (UCI, EME), Isla Cerralvo, 24°14'N- 109o51'W, IV- 16- 1962, R. Moran. Diagnosis. — Eusattus ceralboensis is most similar to E. catalinensis Doyen, differing in having much more coarsely sculptured elytra and a distinct fringe of hypomeral setae. Eusattus cienegus, new species (Figure 72) Moderately convex, elongate oval or oval bee- tles with irregularly 4-costate elytra usually col- ored by pale accretions. Male. — Head and pronotum with duplex punctation, large punctures about 1 Vi eye facets in diameter, small punctures V3-V4 that size, often barely visible at 50 x ; frons with large punctures separated by about 2-4 puncture diameters. slightly denser near epistomal suture, obscured on epistomum; epistomum shallowly, narrowly, and angulately emarginate (as in Fig. 60); medial epistomal suture faint, lateral sutures less ob- scured. Antennal segment length ratios as fol- lows: 3.0:1.0:1.6:1.6:1.5:1.7:1.6:1.6:1.5:1.4:1.8; apical segment about 1 . 1—1.2 x longer than broad, acutely rounded or right-angled; scape and fla- gellum set with short, appressed setae. Mentum about 1.5 x broader than long, anterior corners rounded, anterior margin shallowly to moder- ately emarginate; submentum absent or obsolete; gula narrowed anteriorly, about '/5 width of men- tum. Pronotum 2.0-2.4 x broader than long; disk with coarse punctures shallow, often obscured, separated by 2-3 puncture diameters, becoming denser anterolaterally; hypomeron glabrous, shining, somewhat opaque, with wispy fringe of submarginal setae not reaching pronotal margin; 50 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 sternum punctato-granulose, sparsely set with short, appressed setae; prosternal process 1.2- 1.4 x broader behind than between coxae, un- margined, sparsely punctate. Elytra with lateral margins subparallel in an- terior half, then evenly convergent to apex; disk bearing 2 crenulate, low, rounded, and widely separated costae on each side; intercostal depres- sions irregularly raised into sinuous, anastomos- ing bosses, these more or less coalesced to form 2-3 additional, poorly defined costae, depres- sions filled with pale accreted material, produc- ing distinctive, contrasting pattern; pseudepi- pleural margin explanate (as in Fig. 40); pseudepipleuron shallowly concave, rugulose; epipleuron alutaceous. Metasternum moderately coarsely punctate, with few short setae laterally; abdominal sternites with duplex punctation. Profemur and mesofemur sparsely set with very short, bristle-like setae; protibia with attenuate apical process extending % distance to apex of basal tarsomere; outer margin with irregular row of spinules extending xli-lh distance to apex, sep- arated by 1-4 spinule widths (as in Fig. 83); tar- somere length ratios as follows: 4.4:1.1:1.0:0.9: 3.4 (pro-); 5.8:2.2: 1 .9: 1 .4:3.9 (meso-); 8.4:3. 1 :2.6: 4.8 (meta-). Female. — Apical protibial process extending to apex of basal tarsomere; coxite length ratio to paraproct length = 0.48. Measurements. — EL 8.0-9.1 mm; EW 6.6- 7.5 mm; PL 3.0-5.3 mm; PW 6.2-7.3 mm; BD 4.3-5.0 mm. Holotype Male (CAS) and 9 paratypes (RLA and EME) from Mexico, Coahuila, 20.9 km S Cuatro Cienegas, gypsum dunes, X-l-1976, M. E. Mispagel; 3 paratypes (EME), 9 mi SW Cuatro Cienegas, V-29-1981, J. Doyen and J. Liebherr. Diagnosis. —Eusattus cienegus is similar to E. pons Triplehorn, but differs markedly in elytral sculpturing and has the dorsum smoothly convex (pronotum strongly tumid in pons). It is similar to reticulatus (Say), differing in its more elongate, subparallel body form (broadly ovoid in reticu- latus and elytral sculpturing). In cienegus, the gula is greatly narrowed anteriorly (Fig. 29) and the submentum is obsolescent; in reticulatus, the gula is slightly narrowed and the submentum is a distinct, rectangular sclerite. Distribution and Habitat. — Restricted to the Cuatro Cienegas Basin in Coahuila, Mexico. I collected specimens (adults and larvae) from about the bases of woody shrubs bordering the gypsum dunes SW of Cuatro Cienegas. Exca- vation of shrubs on the dunes, pitfall trapping, and nocturnal searching failed to produce spec- imens. Although some individuals may wander onto adjacent dunes, E. cienegus lacks the mor- phological specializations characteristic of sand- dwelling species, and probably crawls about in the litter collected about low-growing shrubs. This habitat is common to several members of the reticulatus species group. Additional Material Examined. — Mexico: Coahuila: 9 mi SW Cuatro Cienegas. V-29-81, J. Doyen and J. Liebherr (7 corpses). Eusattus costatus Horn Eusattus costatus Horn. 1870:293. pi. 15: 1883:304: 1894: 348, 423 (key). Megasattus costatus, Casey 1908:63; Blaisdell 1943:191. Megasattus sternalis Blaisdell, 1923:268; 1943: 191, new syn- onymy. Convex, broadly oval, black beetles with ru- gose to scabrous, usually costulate elytral sculp- turing. Male. — Head and pronotum with duplex punctation; frons set with large punctures about 1-2 eye facets in diameter, separated by about 2-3 puncture diameters, sometimes becoming denser on epistomum; small punctures V4-V6 that size, separated by 1 puncture diameter or less; epistomal suture slightly impressed or often ob- scured; epistomum narrowly and weakly to mod- erately emarginate; antennal segment length ra- tios as follows: 3.6: 1 .3:2.8:2.2:2.0:2. 1 :2.2: 1 .8: 1.7: 1.6:2.2; terminal segment 1 . 1 — 1.4 x longer than broad; apex acutely angulate; mentum about 1 .4 x broader than long, almost flat, with few scattered setigerous punctures; anterior corners acute, rounded; anterior border moderately to deeply and usually angulately emarginate (Fig. 60); lat- eral borders slightly raised in posterior half; sub- mentum slightly broader than base of mentum; gula anteriorly 0.4-0.7 x width of submentum. Pronotum 1.9-2.4x broader than long, usu- ally widest at base or just before, or occasionally widest just behind middle; disk set with coarser punctures 1-2 eye facets in diameter, separated by 1-4 puncture diameters, sometimes muricate and setigerous near lateral margins: finer punc- tures about 'A-'A that size, separated by 1-2 puncture diameters; hypomeron longitudinally furrowed, especially ventrally, glabrous except DOYEN: SYSTEM ATICS OF EUSATTUS AND CONISA III S 51 for sparse fringe of submarginal setae which pro- ject to margin or slightly beyond; sternum punc- tate to punctato-rugulose; prosternal process punctate or nearly glabrous, margined or un- margined, about 1.5 x wider behind than be- tween coxae. Elytra punctato-rugulose to coarsely rugose or scabrous and usually faintly, sometimes strongly costate; pseudepipleural margin obtuse, round- ed; pseudepipleuron flat, usually bearing a few setae basally. Mesosternum between coxae sub- equal in width to prosternal process, deeply ex- cavate. Metasternum coarsely punctate to punc- tato-rugulose; anterior 3 abdominal sternites finely, sparsely punctate, muricately punctate, or tuberculate. Pro femur with sparse dorsal fringe of long erect setae; tibia with apical process extending to 2nd tarsomere; outer margin with row of spinules extending V:-2/? to apex, spinules subcontiguous basally, separated by 2-4 spine widths apically (as in Fig. 78). Meso- and metafemora with few moderately long hairs dorsally; tarsomere length ratios as follows: 4.4:1.1:1.0:0.9:3.7 (pro-); 5.2:2.2:1.8: 1.5:4.9(meso-); 7.8:2.8:2.2:5.5 (meta-). Female.— Coxite length ratio to paraproct length, 0.33-0.40. Measurements. — EL 7.6-15.0 mm; EW 5.6- 12.0 mm; PL 2.5-4.8 mm; PW 5.3-10.9 mm; BD 4.4-8.8 mm. Holotype Female. — Horn Collection, MCZ. Type Localities.— E. costatus, Baja Califor- nia; sternalis, Angeles Bay, Baja California Norte. Diagnosis.— Eusattus costatus displays con- siderable geographic variation in elytral sculp- turing. Individuals from the northern half of the peninsula have rugose elytra similar to those of E. catavinus Doyen. E. catavinus differs in its much stouter hindbody (0.86 < EW/EL < 0.95; in costatus 0.72 < EW/EL < 0.79). The elytra are more coarsely sculptured in catavinus, and the paraprocts of the ovipositor much shorter. Individuals of costatus from the southern half of the peninsula have scabrous, usually costulate elytra. Sculpturing becomes gradually finer along a north-south trend from the vicinity of Comon- du to the Cape region. Finely sculptured indi- viduals are similar to the most coarsely sculp- tured individuals of E. planulus Doyen, which differs in the more elongate, parallel-sided hind- body (0.67 < EW/EL < 0.75). E. aridus Doyen Figure 73. Known distribution of Eusattus costatus. is similar to E. costatus, differing in the sharply attenuated aedeagus (Fig. 51) (bluntly rounded in costatus). E. erosus is superficially similar to costatus, but has rounded rugae on the elytra, and nearly lacks the lateral fringe of setae on the hypomeron. The holotype resembles specimens from the vicinity of Comondu, Baja California Sur, and the two syntypes are from Isla Santa Margarita and San Luis [Gonzaga], on the Magdalena Plain south of Comondu. Distribution (Fig. 73) and Habitat.— Eu- sattus costatus formerly ranged from San Diego Co., Calif., south to Cabo San Lucas. Recent col- lections have been made no further north than Colonia Guerrero, about 140 miles south of the border. The beetles occur in a variety of arid habitats, including coastal scrub, chaparral, and riparian woodland in the north, and desert scrub 52 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 and thorn forest in the south, at elevations below 600 m. Additional Material Examined. — California: San Diego Co. (2); southern California ( 1 ): Mexico: Baja California Norte: Johnson Ranch [nr. Colonet], IV- 19- 1935 (1); 10 mi NE Co- lonia Guerrero, Rio Santo Domingo, 111-29/30-69 (1); 5.7 mi E Hamilton Ranch, Rio Santo Domingo, IV-24-1963 (10); 8 mi N San Quintin, VI-31-73 (1); San Quintin, V- 13- 1938 (1); vie. San Quintin, V-26-55 (3); Santa Maria Beach, Bahia San Quintin, 111-19-1972 (1); 1 mi N El Socorro, VII-14-1979 (1); Mission Calamajue, IV- 16- 1962 (2); 2 mi W Mission Cala- majue, VI-2-1981 (6); El Desengano [nr. Punta Prieta], VI- 15- 1970 (1); 31.7 mi W Bahia de Los Angeles, VII-12-1979 (1); Bahia de Los Angeles, V- 1965 (1); 1 mi S Bahia de los Angeles, VI-1 1-1967 (1); 2 mi S Los Angeles Bay, 111-19-1951 (1); 14 mi S Rosarito, V-5-1977 (4); Miller's Landing, IV-6-1976 (1). Baja California Sur: 2 mi E San Ignacio, VII-6-1979 (1); 25 mi WSW San Angel, I V-5- 1 96 1 ( 1 ); Mulege, X-26- 1 965 ( 1 ); 3 km W Loreto, XI-26-77 (6); La Purisima, X-1923 (3); Comon- du, V-24-1947 (1); 19 mi SW Comondu, VI-23-1967 (1); San Jorge, XI- 1 5- 1 968 (2); VI- 1 6-73 ( 1 ); Pozo Grande, X-24- 1 968 (1); Santo Domingo, XI- 16- 1941 (1); Puerto San Carlos, VI- 15/16-1973 (1); 1 mi S Palo Blanco, 1400', XI-8-1968 (5); 2 mi SE Santa Rita, 1 000', XI- 1 6- 1 968 (2); 1 3 mi N Villa Con- stitucion, 1 000', XI-7- 1 968 (25); 3.3 mi S El Cien, IX-26- 1981 (1); San Antonio, 1300', X-9- 1968 (1); La Burrera, 12miENE Todos Santos, 1700', XII- 19- 1979 (4); 4.9 mi E Mex. 1 on Hutamonte-San Jose Rd., X-10-1981 (1); San Carlos, IX-25- 1981 (1). Eusattus erosus Horn Eusattus erosus Horn, 1870:294, pi. 15. Megasattus erosus, Casey 1908:63. Convex, broadly oval, black beetles with ru- gose or rugulose elytra. Male. — Head with duplex punctation; frons set with large punctures about 1-3 eye facets in diameter, separated by about 1-3 puncture di- ameters, becoming denser above eyes and coarser and denser near epistomal suture; small punc- tures about V4-V6 that size, separated by about 1 puncture diameter. Epistomal suture faint, fre- quently obscured by cranial sculpturing; episto- mum shallowly to deeply emarginate (Figs. 60, 61); antennal segment length ratios as follows: 3.0:1.4:2.7:2.0:2.0:2.1:2.0:1.9:1.7:1.5:2.3; ter- minal segment about 1.2 x longer than broad; mentum about 1.4 x broader than long, almost flat, glabrous or subglabrous, anterior corners acute, broadly rounded; anterior border deeply, evenly or angulately emarginate; submentum slightly wider than base of mentum; gula ante- riorly about xh width of submentum. Pronotum 2.2-2.5 x broader than long; disk with simple or duplex punctation; large punc- tures about as large as eye facets, separated by 2- 6 puncture diameters, denser laterally and an- teriorly; hypomeron glabrous except for wispy submarginal fringe of setae that do not usually reach pronotal margin; sternum weakly rugulose, glabrous or sparsely setose; prosternal process punctate or rugulose anteriorly, becoming sub- glabrous posteriorly. Elytral disk moderately rugose, noncostate; pseudepipleural margin slightly obtuse to slightly acute; pseudepipleuron flat or feebly concave, weakly to strongly rugose; epipleuron with a few fine punctures, nearly glabrous; metasternum finely to moderately coarsely punctate medially, becoming muricately so laterally; laterally setose or not; anterior 3 abdominal sternites with fine, sparse, simple or duplex punctation. Profemur and mesofemur with sparse dorsal fringe of recumbent setae; protibia with apical process extending to apex of basal tarsomere (as in Fig. 78); outer margin with row of spinules extending about % distance to apex, separated by about 1 spinule width; tarsomere length ratios as follows: 4.3: 1.4: 1.5: 1.8:5.2 (pro-); 7.9:3.7:2.4: 1.9:6.0 (meso-); 11.4:4.2:3.2:7.5 (meta-). Female.— Coxite length ratio to paraproct length a 0.45. Holotype Male (MCZ) from Baja California (Wm. Gabb). Diagnosis.— Eusattus erosus is most similar to catavinus Doyen. In erosus, the elytra are more finely rugose, the smaller punctures on the pro- notal disk are usually so minute as to be almost invisible at 50 x , and the fringe of setae on the hypomeron is either absent or very spare, seldom reaching the pronotal margin. In catavinus, the elytra are much more coarsely sculptured, the pronotal punctation is clearly duplex, with spars- er large and denser fine punctures, and the fringe of setae is much denser, extending beyond the pronotal margin and visible from above. The elytra are relatively shorter and broader in ca- tavinus (0.79 < EW/EL < 0.88) than in erosus (0.73 < EW/EL < 0.81). Habitat.— E. erosus manuelis was collected beneath the canopies of shrubby vegetation near a brackish water lagoon on Isla Espiritu Santo, and numerous corpses were found on sand dunes on Isla Partida. Variation.— E. erosus occurs in several weak- ly differentiated but strongly disjunct popula- tions in Baja California. This variation is rec- ognized here at the subspecific level. DOYEN: SYSTEM ATICS OF EUSATTUS AND CON1SATTUS 53 Key to the Subspecies of Eusattus erosus 1 . Elytra with pseudepipleural margins slight- ly obtuse 2 Elytra with pseudepipleural margins slight- ly acute erosus laeviventris 2. Pronotum with duplex punctation; hypo- meron with sparse fringe of submarginal setae reaching margin erosus erosus Pronotum with apparently simple puncta- tion, hypomeron glabrous or with few short setae that do not reach margin erosus manuelis Eusattus erosus erosus Horn Eusattus erosus Horn, 1870:294, pi. 15; 1894:349, 423 (key). Megasattus erosus, Casey 1908:63; Blaisdell 1923:265 (in part); 1943:190 (in part). Pronotal disk with duplex punctation, smaller punctures easily visible at 50 x ; hypomeron with sparse fringe of marginal setae barely reaching pronotal border; elytral disk and pseudepipleu- ron strongly rugose; pseudepipleural margin ob- tuse; abdominal sternites with duplex puncta- tion. Measurements. — EL 11.7-12.1 mm;EW9.1- 9.6 mm; PL 3.6-3.7 mm; PW 8.6-9.0 mm; BD 6.5-7.0 mm. Holotype Male (MCZ) from California (Wm. Gabb). Diagnosis.— E. erosus erosus is distinguished by its duplex cranial and abdominal punctation from E. erosus manuelis and E. erosus laeviven- tris. Additional Material Examined (Fig. 74). — Mexico: Baja California Sur: Comondu, 11-23 (USNM: 1); VII-1953 (USNM: 1). Eusattus erosus manuelis (Blaisdell) Megasattus erosus manuelis Blaisdell, 1923:266; 1943:190; Doyen, 1972:369 (morphology). Pronotal disk with apparently simple punc- tation (smaller punctures exceedingly minute, in- visible at 50 x ); hypomeron entirely glabrous or with very wispy fringe of few submarginal setae that do not reach pronotal border; elytral disk moderately rugose; pseudepipleuron weakly ru- gose; pseudepipleural margin obtuse; abdominal sternites with apparently simple punctation. Measurements. — EL 9.1-14.8 mm; EW 6.9- 1 1.7 mm; PL 2.9-4. 1 mm; PW 6.5-9.7 mm; BD 4.9-7.7 mm. Holotype Male (CAS) from Mexico, Baja \ . •) \ A. a { ) ■ ( S ■ H j c. k„^ £>- — . - 1 \ >i u\* ^-J \ \ ° V }' M \ l %J> K \ <• "* 4k Y> 0 EROSUS MANUELIS ^v \ O. 5\ (J E LAEVIVENTRIS \ O E EROSUS N. ■ CATAVINUS \ Figure 74. catavinus. Known distribution of Eusattus erosus and E. California Sur, Isla Espiritu Santo, V-31-1921, J. R. Slevin. Diagnosis.— E. erosus manuelis is distin- guished from E. erosus erosus by the simple cra- nial and abdominal punctation. It differs from E. erosus laeviventris in having the pseudepi- pleural margin slightly obtuse. Its distribution is restricted to the vicinity of La Paz Bay and Isla Espiritu Santo. Additional Material Examined (Fig. 74). — Mexico: Baja California Sur: El Crucero, VII-3 1 - 1 968 ( 1 ); 1 2.4 mi E La Pa? on road to Las Cruces, XII-23-58 (9); Arroyo Saltito nr. Las Cruces, 1-23-59 (1); Isla Espiritu Santo, 11-22-1953 (1); Isla Espiritu Santo, Bahia San Gabriel, HI-3 1-1974 (28); Isla Par- tida, 111-23-1953 (5), IV-12-1974 (17). Eusattus erosus laeviventris (Blaisdell) Megasattus laeviventris Blaisdell, 1923:167; 1943:192. Pronotal disk with apparently simple punc- tation (smaller punctures exceedingly minute, in- 54 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 visible at 50 x); hypomeron with very sparse fringe of very short setae, not approaching pro- notal border; elytral disk moderately rugose; pseudepipleural margin acutely rounded; ab- dominal sternites with apparently simple punc- tation. Measurements. — EL 13.9-14.8 mm; EW 10.9-11.5 mm; PL 4.3-4.4 mm; PW 9.8-10.5 mm; BD 7.5-8.0 mm. Holotype Female (CAS) from Mexico, Baja California Sur, Isla Santa Cruz, VI- 1 1-1921, I. M. Johnson. Diagnosis.— E. e. laeviventris is distinguished from the other subspecies of E. erosus by its acutely rounded pseudepipleural margins. Bahia Escondido (see below) is on the main- land near the south end of Isla del Carmen. A single specimen collected by J. P. Figg-Hoblyn and labeled Isla Santa Catalina clearly represents E. e. laeviventris, rather than the very distinctive E. catalinensis (see discussion under E. catali- nensis). Additional Material Examined (Fig. 74). — Mexico: Baja California Sur: Isla Santa Cruz, 111-26- 1 953 (2); Escondido Bay, VI-14-1921 (1). Eusattus franciscanus, new species Convex, broadly oval, black beetles with ru- gulose, faintly carinulate elytra. Male. — Head with duplex punctation; frons set with large deep punctures about l'/2-3 eye facets in diameter, separated by about 1-3 punc- ture diameters, becoming denser, often nearly confluent around epistomal suture; small punc- tures about lA that size, separated by about 1 puncture diameter. Epistomal suture faint, ob- scured or obliterated medially by cranial sculp- turing; epistomum weakly to moderately and evenly emarginate (as in Fig. 60). Antennal seg- ment length ratios as follows: 3.5:1.3:2. 1 : 1.5:1.8: 1.6:1.4:1.5:1.4:1.3:1 .6; terminal segment slightly broader than long, apex right-angled or slightly obtuse; mentum about 1.3 x broader than long, sparsely set with long setae; anterior corners acute, broadly rounded; anterior border deeply, evenly emarginate; submentum slightly broader than base of mentum; gula anteriorly about xh width of submentum. Pronotum about 2.2-2.4 x broader than long; disk with apparently simple punctation, or with fine punctures barely visible at 50 x; large punc- tures about 2 eye facets in diameter, deep, coarser and denser laterally and anteriorly; hypomeron glabrous except for sparse marginal fringe of se- tae extending just beyond pronotal margin; ster- num rugulose, sparsely set with semierect, mod- erately long setae; prosternal process unmargined, punctate anteriorly, becoming nearly glabrous posteriorly. Elytral width at base subequal to pronotal width; disk set with numerous irregular, shallow erosions producing a rugulose appearance; each elytron usually with 4-5 faint but evident ca- rinulae nearly reaching apex; pseudepipleural margin right-angled in cross-section, broadly rounded (as in Fig. 37); pseudepipleuron flat or feebly concave, rugulose; metasternum moder- ately coarsely punctate, sparsely setose; anterior 3 abdominal sternites with apparently simple, sparse punctation, or with duplex punctation, fine punctures barely visible at 50 x . Profemur and mesofemur with sparse dorsal fringe of moderately long, inclined or retrorse setae; protibia with bluntly rounded apical pro- cess extending about to apex of basal tarsomere (as in Fig. 78); outer margin with irregular row of spinules extending about xh distance to apex, separated by about 1-4 spinule widths; tarso- mere length ratios as follows: 4.0: 1 .4: 1 .0: 1 .0:4.0 (pro-); 5.8:2.0:1.8:1.4:4.2 (meso-); 8.2:3.0:2.2:4.5 (meta-). Female.— Coxite length ratio to paraproct length =s 0.45. Measurements. — EL 7.7-9.9 mm; EW 6.3- 8.0 mm; PL 2.4-3.1 mm; PW 5.4-7.0 mm; BD 4.2-5.3 mm. Holotype Female (CAS) and 12 paratypes from Mexico, Baja California Sur, Isla San Fran- cisco, 1-7-1976, T. M. Glimme. Diagnosis.— Eusattus franciscanus is most similar to E. erosus manuelis Blaisdell, but ma- nuelis is considerably larger, has much coarser elytral sculpturing, and has relatively narrower elytra (0.73 < EL/EW < 0. 8 \)lhan franciscanus (0.79 < EL/EW < 0.88). The profemoral setae are much longer in franciscanus than in ma- nuelis. Discussion. — Two damaged specimens col- lected by J. P. Figg-Hoblyn (CAS) and labeled Isla San Francisco and Isla San Jose are outside the size range listed for E. franciscanus and have narrower elytra. These specimens probably came from another gulf island. Another specimen col- lected by Figg-Hoblyn and labeled Santa Cata- DOYEN: SYSTEMATICA OF EUSATTUS AND CON1SAI 1 1 S 55 lina Island clearly originated elsewhere (see ac- counts for E. catalinensis and E. erosus laeviventris), suggesting that locality data for his 1953 trip were confused. Habitat. — The type series was collected from maritime dunes. Corpses were taken from be- neath stones in strongly maritime situations. Additional Material Examined. — Mexico: Baja Califor- nia Sur: Isla San Francisco, IV-9-1952 (1). 111-24-1953 (1), I V-9- 1955 (2); unnamed cove, S end, coastal dunes, IV- 1 - 1 974 (10 corpses); Isla San Jose, Bahia Amortajada. IV- 1-1974 (2); Puma Ostiones, IV-7-1974 (1). Eusattus laevis LeConte (Figure 75) Eusattus laevis LeConte, 1866:113; Horn 1870:294; 1883: 205; 1894:349, 423 (key). Eusattodes laevis, Casey 1908:64; Blaisdell 1943:192. Convex, pyriform or oval black beetles with glabrous, weakly shining cuticle. Male. — Head with duplex punctation. Frons set with coarse punctures about V3-V2 eye facet in diameter, separated by about 3-8 puncture diameters, becoming denser near epistomal su- ture; fine punctures about lA eye facet in diam- eter or less, often barely visible at 50 x , separated by 1-2 puncture diameters; epistomum with coarse punctures separated by 2-3 puncture di- ameters; anterior border narrowly and moder- ately to deeply emarginate medially (as in Fig. 59), slightly indented at lateral sutures; epistomal suture fine, weakly impressed, sometimes ob- scured medially. Antennal segment ratios as fol- lows: 3.3:1.3:2.2:1.9:1.8:1.8:1.7:1.6:1.6:1.5:1.8; apical segment about 1 . 1 x longer than broad, apex rounded or broadly right-angled; mentum about 1 .3-1.4 x broader than long, flat, glabrous; anterior corners broadly rounded, anterior bor- der moderately to deeply, and arcuately emar- ginate; submentum rectangular, as wide as men- tum at base; gula about lh width of mentum at base. Pronotum 2. 1-2.4 x broader than long; disk set with punctures about lA-Vs eye facet in di- ameter, separated by about 2-6 puncture di- ameters; hypomeron with an exceedingly wispy submarginal fringe of short setae, often appearing completely glabrous; sternum very shallowly punctato-rugulose, sparsely set with short, de- cumbent setae; prosternal process unmargined, with a few fine punctures, 1 .4-1 .7 x broader be- hind than between coxae. Elytra widest behind middle; disk alutaceous, Figure 75. Eusattus laevis LeConte. sparsely set with exceedingly fine punctures, barely visible at 50 x; pseudepipleural margin broadly rounded, undefined; epipleuron aluta- ceous, glabrous; mesosternum between coxae 0.5- 1.0 x width of prosternal process; metasternum finely, sparsely punctate, with a few decumbent setae laterally; abdominal sternites sparsely and exceedingly finely punctate. Profemur and mesofemur sparsely set with short decumbent setae; protibia with rounded apical process extending to apex of basal tarso- mere; outer margin with row of spinules extend- ing V2— 2A distance to apex, separated by about 1- 3 spinule widths (Fig. 80); tarsomere length ratios as follows: 5.3:1.7:1.6:1.6:4.3 (pro-); 8.0:3.4:3.2: 2.3:5.4 (meso-); 10.6:4.3:3.4:6.2 (meta-). Female.— Coxite length ratio to paraproct length a 0.50. Measurements. — EL 9.0-13.3 mm; EW 6.9- 10.3 mm; PL 3.0-4.0 mm; PW 6.2-8.5 mm; BD 4.8-7.4 mm. Lectotype Female (MCZ) from Mexico, Baja California Sur, Cabo San Lucas. Diagnosis.— Eusattus laevis differs from all other members of the genus except E. reticulatus (Say) in the extremely fine sculpturing of its pronotum and elytra, and in being practically devoid of setae; reticulatus is readily distin- 56 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 Figure 76. Known distribution of Eusattus laevis in south- ern Baja California. guished by its explanate pseudepipleural margin (rounded in laevis). E. laevis is superficially sim- ilar to E. rudei Doyen, but the epipleuron of rudei is gradually broadened, and the venter is densely set with long, erect setae. Distribution (Fig. 76) and Habitat. -E. lae- vis occurs in a variety of habitats, including Cape thorn forest, riparian associations, and open grassland, at elevations ranging from sea level to over 1800 m. The great majority of specimens are from the Cape region from La Paz south to Cabo San Lucas, and from Isla Ceralbo. Outside this area, single collections have been made at San Domingo (=Santo Domingo?) and Santa Ro- salia, about 220 and 420 km north of La Paz respectively. Possibly these labels are in error; a single specimen labeled San Felipe is certainly in error. Eusattus mexicanus Champion (Figure 77) Eusattus mexicanus Champion, 1892:510. Strongly convex, very broadly oval, black or blackish-brown beetles, with reticulate elytra usually colored by accreted material. Male.— Head and pronotum with punctation duplex or apparently simple, with fine punctures barely visible at 50 x ; frons set with coarse punc- tures about 2-3 eye facets in diameter, coarsest posteriorly along vertex, separated by about 2- 4 puncture diameters, sometimes with larger, impunctate areas, becoming denser along epi- stomal suture; epistomum shallowly, evenly emarginate medially, deeply indented at lateral sutures (as in Fig. 60); epistomal suture distinct, weakly impressed. Antennal segment length ra- tios as follows; 3.8:1 .6:2.3: 1.7:1.6: 1.8: 1 .8: 1.9:1.9: 1 .7:2.0; terminal segment about as broad as long, with acute apex; mentum coarsely punctate, cres- centic; about 1.7-1.9 x broader than long, with paired foveae near posterior corners; anterior corners broadly rounded; anterior border mod- erately and evenly emarginate; submentum slightly broader than base of mentum; gula an- teriorly about Vi width of submentum. Pronotum about 1.9-2.1 x broader than long; lateral margin evenly arcuate or weakly angulate near middle, and nearly straight anteriorly and posteriorly, slightly divergent in posterior fifth; disk set with shallow, nearly obsolete punctures about size of eye facets, separated by about 2-4 puncture diameters; sparse punctures, barely vis- ible at 50 x, sometimes apparent; hypomeron polished, with sparse submarginal fringe of setae not reaching pronotal border; sternum punctato- rugulose, sparsely set with short, appressed setae; prosternal process about 1.75 x broader behind than between coxae, unmargined, punctate. Elytra with lateral margins arcuate, more abruptly so posteriorly; disk on each side with 3 irregular, slightly raised, rounded and polished ridges, these connected by anastomosing, trans- verse ridges; depressions between ridges filled with accreted material, often of contrasting hue, producing distinctive reticulate appearance; pseudepipleural margin obtusely rounded (as in Fig. 37); pseudepipleuron flat, sparsely and ir- regularly set with broad, shallow punctures, these within shallow excavations, producing an eroded appearance; epipleuron abruptly broadened from about middle of 1st sternite to humerus; meta- sternum sparsely, finely punctate to almost gla- brous; abdominal sternites coriaceous, finely, ob- soletely punctate. Profemur and mesofemur set with short, de- cumbent setae; protibia with spatulate, distally rounded process extending to apex of 1st or 2nd DOYEN: SYSTEM AT1CS OF EUSATTUS AND CONISATTUS 57 Figure 77. Eusattus mexicanus Champion. tarsomere; outer margin with row of spinules extending xk-2h distance to apex, separated by about 1-3 spinule widths (Fig. 81); tarsomere length ratios as follows: 4.5:1.6:1.6:1.5:5.1 (pro-); 6.0:2.0:2.0:1.7:5.5 (meso-); 8.5:3.5:1.9:5.9 (meta-). Female. —Apical process of protibia extending to apex of 3rd or 4th tarsomere (Fig. 81); coxite length ratio to paraproct length = 0.50. Measurements. — EL 7.2-1 1.4 mm; EW 6.7- 9.3 mm; PL 2.6-3.9 mm; PW 5.2-8.1 mm; BD 4.2-6.4 mm. Lectotype Male (?) (BMNH) hereby desig- nated, from Mexico, Jalisco, Sayula; 7 syntypes (1 AMNH) same data; 3 syntypes from Colima, Colima City; 1 syntype from Durango, villa Ler- do; 1 syntype from Guerrero, Chilpincingo. Diagnosis.— Eusattus mexicanus is similar to E. venosus Champion (see diagnosis for venosus). It is similar to E. pons Triplehorn, differing in the rounded pseudepipleural margin (narrowly explanate in pons), in the reticulate elytra (coarsely rugose in pons), and in the transverse, crescentic mentum (broader, trapezoidal in pons). Distribution (Fig. 87).— Jalisco and Colima, Mexico. Additional Material Examined. — Mexico: Jalisco: mtns. N Ajijic, 7500', IX-20-1964 (9); arroyo N Ajijic, 5250', VII- 1-1964 (1), VII-29-1964 (1); 4 mi N Atenquique, XI-25-1950, (2); Atenquique, XII-5- 1 948 ( 1 ); envir. Guadalajara, 1901(1); La Floresta, Lago de Chapala, 1510m, IX-4/5- 1 977 (2); Ocot- lan (1); Sayula, XI-23 (14); VII-30-1964 (1). Colima: no ad- ditional data (2); X-1954 (1); Volcano Mex (2). Eusattus planulus, new species Eusattus productus. Horn 1894:349 (in part); Blaisdell 1943: 194 (in part). Convex, elongate, ovoid black beetles with alutaceous or finely rugulose elytra. Male. — Head and pronotum with duplex punctation; frons with large punctures about 1 Vi eye facets in diameter, separated by about 1-3 puncture diameters, slightly denser near episto- mal suture; small punctures about Vb that size, 58 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figures 78-83. Foretibiae. posterior view. 78. Eusattus andus. 79. E. cienegus. 80. E. laevis. 81. E. mexicanus. 82. E. pons. 83. E. reticulatus. separated by 1 puncture diameter or less; medial epistomal suture weakly impressed; epistomum shallowly, usually angulately emarginate; anten- nal segment length ratios, as follows: 3.7: 1 .5:2.8: 2.2:2.0:2.0:2.0: 1 .8: 1 .7: 1 .5:2.2; terminal segment 1 . 1 — 1 .2 x longer than broad; apex acutely an- gulate; mentum almost flat, sparsely setose; an- terior corners acute, narrowly rounded; anterior border deeply, usually angulately emarginate; submentum slightly broader than base of men- turn; gula anteriorly about 0.4 x width of sub- mentum. Pronotum 2.1-2.4x broader than long, disk set with punctures slightly larger than eye facets, separated by 2-5 puncture diameters, becoming slightly denser anterolaterally; hypomeron lon- gitudinally furrowed, glabrous except for sub- marginal fringe of setae which project beyond lateral margins; sternum punctato-rugulose, fine- ly, sparsely setose; prosternal process unmar- gined, nearly glabrous, sparsely punctate, 1.5 x wider behind than between coxae. Elytra shining, alutaceous to finely rugulose, sometimes very faintly costulate; lateral elytral margin obtuse, rounded; pseudepipleuron flat; epipleuron finely, sparsely punctate, with a few setae basally; metasternum coarsely punctate; anterior 3 sternites finely, sparsely punctate. Profemur with sparse dorsal fringe of long, erect setae; tibia with apical process extending to 2nd or 3rd tarsomere, outer margin with row of spi- nules extending Vi-2h distance to apex; spinules subcontiguous basally, separated by 3-4 spine widths apically, Mesofemur with dorsal fringe of erect, moderately long setae; metafemur sub- glabrous; tarsomere length ratios as follows: 4.3: DOYEN: SYSTEM ATICS OF EUSATTUS AND CONISATTUS 59 1.1:1.0:0.9:4.5 (pro-); 6.2:2.4:2.0:1.6:5.0 (meso-); 8.4:3.0:2.4:5.0 (meta-). Female. -Coxite length ratio to paraproct length a 0.33. Measurements. -EL 10.0-12.4 mm; EW 7.0- 8.7 mm; PL 3.0-3.8 mm; PW 7.0-8.3 mm; BD 5.3-6.7 mm. Holotype Male (CAS) Mexico, Baja Califor- nia Sur, La Paz, 40 m (Cape Thorn Forest), XII- 23-1974, E. L. Sleeper. Paratypes, La Paz, XII- 19-1974, W. W. Middlekauff ( 1 , EME); La Paz, XII-19-1973, W. W. Middlekauff(l, EME); 4 mi E La Paz on road to Los Cruces, XII-23-1958, H. B. Leech (1, CAS); 20 km E Colonia de la Toba, 400', XI-23- 1 968, E. L. Sleeper ( 1 , EME); El Pescadero, IV- 16- 1979, M. Wasbauer (1, CDFA); 4 mi S El Pescadero, X-24-1968, E. L. Sleeper and J. Moore (1, CSULB); Playa Lobos, nr. Todos Santos, XII- 10- 1979, J. Doyen and M. Wasbauer ( 1 , EME); 2 mi S Todos Santos, X- 1 6- 1968, 100', E. L. Sleeper and J. Moore (1, CSULB). Diagnosis. — Coarsely sculptured individuals of Eusattus planulus are very similar to finely sculptured specimens of E. costatus, but the hindbody of planulus is more elongate (0.72 < EW/EL < 0.79 for costatus; 0.67 < EW/EL < 0.75 for planulus). The elytra of planulus are usually more parallel-sided than in costatus. Smooth individuals of planulus resemble E. ru- dei Doyen and E. productus LeConte. In pro- duces, the elytra are finely tuberculate (punctate or rugulose in planulus), the epipleura are dense- ly setose anteriorly (few scattered setae in plan- ulus), and the spine row extends the entire length of the outer foretibial margin (Vi-2/) its length in planulus). In rudei. the elytral punctures are very fine (subequal to an eye facet), the body is more strongly convex, and the outer margin of the fore- tibia is nearly straight (Fig. 133). Distribution (Fig. 84) and Habitat.— Thorn- forest habitats below 500 m in the Cape region of Baja California. Additional Material Examined. — Mexico: Baja Califor- nia Sur: 14 mi W La Paz, X-23/24-1972 (1); 18 mi W La Paz. XI-25-1968 (2); 19.2 mi W La Paz, XII-31-1958 (1); 27 mi W La Paz. XI-18-1968 (25). Eusattus pons Triplehorn Eusattus erosus. Champion 1892:509; Pallister 1954:43. Eusattus pons Triplehorn, 1968. Strongly convex, very broadly oval black bee- Figure 84. Known distribution of Eusattus planulus in southern Baja California. ties with coarsely rugulose elytra bearing light- colored accretions. Male. — Head and pronotum with duplex punctation; coarse punctures about as large as eye facets, small punctures about l/4-V2 that size; frons with large and small punctures separated by about 1-2 puncture diameters, becoming denser on epistomum and about eyes; episto- mum moderately and evenly emarginate medi- ally (as in Fig. 60), weakly emarginate at lateral epistomal sutures; epistomal suture slightly im- pressed or obscured by punctation. Antennal seg- ment length ratios as follows: 3.6: 1 .7:2.3: 1 .8: 1.8: 1.8:1.8:1.9:2.0:1.7:2.3; apical segment 1.2-1.3 x longer than broad, apex nearly right-angled or acute; mentum about 1 .6-1 .7 x broader than long, with deep submarginal groove along posterior half of sides; anterior corners broadly rounded, acute; anterior border moderately and angulately emarginate. Submentum absent or obsolete; gu- lar apex lh width of mentum. Pronotum about 2.0-2.2 x broader than long, strongly convex, tumid, in lateral silhouette (Fig. 49), broadest at base; disk set with coarse punc- tures separated by 2-4 puncture diameters, be- coming coarser, closer anterolaterally; small punctures separated by 1-2 diameters; hypom- eron opaque, with wispy submarginal fringe of setae not reaching lateral pronotal border; ster- num punctato-rugulose, sparsely set with short, appressed setae; prosternal process about 1.3- 1 .4 x broader behind than between coxae, un- margined, sparsely punctate. Elytra with lateral margins gently arcuate in anterior half, then abruptly convergent to apex; 60 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figure 85. Eusattus reticulatus (Say). disk coarsely, deeply rugose, ridges polished, gla- brous, depressions granulate, usually containing light-colored accretion; producing distinct, con- trasting pattern; pseudepipleural margin expla- nate, crenulate; pseudepipleuron shallowly con- cave, rugose; epipleuron alutaceous; metasternum moderately coarsely punctate, with few setae lat- erally; abdominal sternites with duplex puncta- tion. Profemur and mesofemur sparsely set with very short, decumbent setae; protibia with attenuate apical process extending to apex of 1st or 2nd tarsomere (Fig. 82); outer margin with row of spinules extending Va distance to apex, separated by 1-4 spinule widths; tarsomere length ratios as follows: 4.4: 1 .3: 1 .3: 1 .2:4.0 (pro-); 7.0:2.5:2.4: 2.0:4.7 (meso-); 9.9:3.4:3.0:5.5 (meta-). Female.— Coxite length ratio to paraproct length a 0.40. Measurements. — EL 8.8-10.5 mm; EW 7.3- 10.1 mm; PL 3.3-4.2 mm; PW 6.6-9.2 mm; BD 4.8-6.5 mm. Holotype Male and 3 paratypes (OSUC) from Texas, Brewster County, Green Valley, VII- 14, H. A. Wenzel; additional paratypes, Chisos Mtns., Texas, VII- 17- 1946, D. J. and J. N. Knull (1); VI-26-1961, D. J. and J. N. Knull (1); VIII-7/8- 1962, C. A. and W. E. Triplehorn (2); VII- 10- 1930, H. M. Smith (1); VII-13-1958, W. F. Barr (1); Glenn Springs, VII- 18- 1930, H. M. Smith (1). Diagnosis.— Eusattus pons is most similar in superficial features to E. catavinus Doyen, dif- fering in its carinate pseudepipleural margin (rounded in catavinus). It is similar to E. venosus Champion, differing in its rugose elytra (reticu- late in venosus), rugose pseudepipleuron (aluta- ceous in venosus), and in having its epipleuron less abruptly broadened at the base. Distribution (Fig. 87) and Habitat.— E. pons occurs in seasonally arid montane and riparian woodland, from 2500-5000 ft elevation. The geographic range extends from the Pecos River in west Texas to the Rio Nazas, Durango, Mex- ico. Additional Material Examined.— Texas: Brewster Co.: Chisos Mtns., VII-12/1 5-1962 (2); Chisos Mtns., Basin, VII- 12-68 (1), VII-28-1937, 5000' (1); E slope Chisos Mtns., VIII- 6-1936, 4800' (1); between Basin Jet. and the Basin, VIII- 12- 1955 (1); Juniper Canyon, VII-20-1928 (1); Window Trail, VIII- 15- 1968 (1); Big Bend National Park, IV-8 to IX- 17 (5); Pecos Co.: Sheffield, VI-30-1948 (1); Pecos River, 6 mi SE Sheffield, VII- 17- 1964(1); Presidio Co.: Presidio, VIII-27-1968 (1); Terrell Co.: 1 mi W Sanderson, 2800', VIII-26-1971 (2). Mexico: Chihuahua: 20 mi SW Camargo, 4500'. VII- 13- 1947 (1); 50 mi S Chihuahua, VII-24-1967 (1); 10 mi N Jimenez, IX- 10- 1 950 ( 1 ). Coahuila: Sierra de los Burros [N of Boquillas], VI- 18- 1958 (2). Durango: Villa Lerdo (7); 3 mi W Villa Lerdo, VIII-24-1946 (1); Tlahualilo, X-20-1935 (12). Eusattus reticulatus (Say) (Figure 85) Zophosis reticulata Say, 1824:250; Gory 1837:187; LeConte 1859a:147. Eusattus reticulatus, LeConte 1851:132; 1858:37; 1859b:14; Lacordaire 1859:221; LeConte 1866:1 12; Horn 1870:293; 1883:304; 1894:423 (key); Champion 1892:509; Cockerell and Fall 1907:202 (distribution); Casey 1924:311; Pal- lister 1954:44; Doyen 1972:369 (morphology); 1977:6 (synonymy); Tschinkel and Doyen 1976:331-335 (biolo- gy)- Discodemus reticulatus, LeConte 1862:223; Casey 1908:61. Discodemus corrosus Casey, 1908:61. Discodemus brevipennis Casey, 1 908:6 1 . Discodemus elongatulus Casey, 1 908:6 1 . Discodemus depressulus Casey, 1908:62. Discodemus subsenceus Casey, 1908:62. Discodemus knausi Casey, 1908:62. Eusattus corrosus, Casey 1924:31 1. DOYEN: SYSTEMATICS OF EUSATTUS AND CONISATTUS 61 Eusattus brevipennis, Casey 1924:311. Eusattus elongatulus, Casey 1924:31 1. Eusattus depressulus, Casey 1924:311. Eusattus subsericeus, Casey 1924:31 1. Eusattus knausi, Casey 1924:31 1. Very broadly oval, moderately convex, black beetles with costate-reticulate elytra. Male. — Head and pronotum with sculpturing usually very shallow, obscured, appearing im- punctate or nearly so, especially on pronotal disk; when well developed, punctation duplex, large punctures about as large as eye facets, separated by about 2-4 puncture diameters on frons; small punctures about Va that size, separated by about 1 puncture diameter; epistomum punctate, fre- quently rugulose or corrugate; anterior border shallowly to moderately and narrowly emargin- ate medially (as in Fig. 60), shallowly concave at lateral sutures; epistomal suture weakly im- pressed, frequently obscured medially. Antennal segment ratios as follows: 3.5: 1 .5:2.5:2.2:2. 1 :2.0: 2.0: 1 .8: 1 .7: 1 .6:2.2; apical segment about 1 .25 x longer than broad, acutely rounded or nearly right-angled; mentum about 1.5 x broader than long, with submarginal groove along posterior V3-V2 of lateral borders; anterior corners broadly rounded; anterior margin moderately to deeply and arcuately emarginate; submentum broader than mentum; gula about Vi width of mentum. Pronotum 2.3-2.6 x broader than long; disk set with shallow punctures about as large as eye facets, separated by about 1-4 puncture diame- ters, more or less obscured, sometimes not ap- parent; hypomeron glabrous except for wispy submarginal fringe of setae not reaching lateral margin; presternum finely punctato-rugulose, sparsely set with short, decumbent setae; pro- sternal process about 1-4 x broader behind than between coxae, sparsely punctate. Elytra with lateral borders arcuate, more strongly so in posterior third; disk with 6 round- ed costae; intercostal depressions raised into irregular, anastomosing transverse rugae, pro- ducing reticulate appearance; depressions occa- sionally filled with light-colored accretion, cre- ating sharply contrasting pattern; pseudepipleural margin explanate, upturned; pseudepipleuron alutaceous, shallowly concave; epipleuron alu- taceous, with few short setae basally; metaster- num sparsely, moderately coarsely punctate, sparsely setose; abdominal sternites with fine, sparse duplex punctation. Profemur and mesofemur sparsely set with short, decumbent setae; protibia with apical pro- cess extending from xh to full length of basal tar- somere; outer margin with row of spinules ex- tending about V4 distance to apex, separated by 1-4 spine widths (Fig. 83). Female.— Coxite length ratio to paraproct length a 0.35-0.40. Measurements. -EL 7.7-13.2 mm; EW 6.7- 11.8 mm; PL 2.5-4.1 mm; PW 6.2-10.5 mm; BD 4.4-7.7 mm. Types. — Holotype not designated. The type material of Zophosis reticulata was destroyed along with the rest of Say's collection, but its identity is clear from his description and the type locality. Type Localities. -Of reticulata, near the Rocky Mountains; brevipennis, the Grand Can- yon, Arizona; corrosus, near El Paso, Texas; de- pressulus, southern Arizona; elongatus, San Ber- nardino Ranch, Cochise Co., Arizona; knausi, Fremont County, Colorado; subsericeus, Arizo- na. Diagnosis. -Eusattus reticulatus is most sim- ilar to E. cienegus Doyen, differing as stated in the diagnosis for that species. It is similar to E. pons Triplehorn, which differs in having the pronotum tumid (streamline in reticulatus) and the elytra coarsely, deeply rugose. E. reticulatus is superficially similar to E. convexus LeConte, which has the epipleuron gradually broadened and the pseudepipleuron rounded (abruptly broadened; explanate in reticulatus). Variation. -Specimens from Colorado are more strongly reticulate with less distinct costae than those from farther south; they correspond to reticulatus (Say) and knausi Casey. Specimens from Yavapai Co., Arizona, have weakly costate, nonreticulate elytra; they correspond to subse- riceus Casey. Habitat and Distribution (Fig. 86). -Sub- arid or seasonally arid grassland, scrub or wood- land associations, ranging from near sea level (lower Colorado River Valley) to about 2500 m are occupied. E. reticulatus is commonest in ri- parian habitats in southern Arizona and New Mexico, where the great majority of specimens have been collected. Adults frequent debris about the bases of shrubs, accumulations of fallen leaves, and similar situations. Collection dates all fall into the period from the last week of June to the first week of October, corresponding to the sum- mer monsoons of the southwestern United States. 62 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 i TZ T- T^ "■ TT: ■.*•;*' " • ' *■•■'■:: \ > ■■ S^rZ' """-' ' ; / .& / ■ | / " ""'■ *, .; :•', :'. - ■>„ I "' T i •** ..--■' ■ -■■ -:. '.■;■■■■■ .../ "I"-*---— -: • ♦ *• :.«S& ■.••* ft •; f -• • .'----- ^ ' A : • ' : ■-■■■■■ r" **-----.--. j Vi*s» * V ';,.- • ;• "" ----_,. o * i •'•':•. :* > h •. *> • •.. -r* ^ < ; • • • '-""-:/-•/-„ 0 <3 , 1 \ I Vl* I V P >V ' '' ^ "\ ( » ) ^ - ' liP \ H kjif * ^ ■ ■■■ ■ ■■■■ ■■■ ■■■-r: ^-:: f Figure 86. Known distribution of Eusattus reticulums. A single record from Kingsville, Kleberg Co., Texas, is not plotted. Eusattus venosus Champion Eusattus venosus Champion, 1892:509. Strongly convex, very broadly oval; black or blackish-brown beetles with weakly carinate pseudepipleural margin and reticulate elytra bearing light-colored accretions. Male. — Head and pronotum with punctation duplex or apparently simple, with fine punctures barely visible at 50 x ; frons set with coarse punc- tures about 1 Vi-2 eye facets in diameter, sepa- rated by 2-3 puncture diameters, or very sparse- ly, irregularly distributed, with large impunctate areas; epistomum slightly more densely punc- tate, especially along suture, to rugosopunctate, shallowly to moderately emarginate medially (as in Fig. 60), slightly indented at lateral sutures; epistomal suture distinct, weakly impressed; an- tennal segment length ratios as follows: 2.7:1.3: 2.0:1.6:1.6:1.8:1.7:1.7:1.8:1.6:1.7; terminal seg- ment about as broad as long, with apex right- angled or slightly acute; mentum about 1 .6-1.7 x broader than long, with irregular submarginal groove along each side becoming deeply im- pressed near base; anterior corners acute, nar- rowly rounded; anterior border moderately and angulately emarginate. Pronotum strongly convex in longitudinal sil- houette, about 1.9-2.1 x broader than long; lat- eral margins evenly arcuate or weakly angulate near middle, and nearly straight anteriorly and posteriorly, slightly divergent in posterior fifth; DOYEN: SYSTEMATICS OF EISA I "ITS AND COS1SA1 TVS 63 disk set with shallow punctures subequal to those on cranium, separated by about 2-4 puncture diameters, usually obscured by accretion; sparse punctures, barely visible at 50 x, sometimes ap- parent; hypomeron opaque, finely granulate, crinkled in coxal region and finely rugulose or tuberculate submarginally; sternum finely punc- tato-rugulose, with scattered, very short, fine, ap- pressed setae; prosternal process about 1 .4-1 .5 x broader behind than between coxae, unmar- gined, sparsely, finely punctate. Elytra with lateral margin arcuate, more abruptly so posteriorly; disk on each side with 2 irregular, slightly raised, rounded and polished ridges, these connected by anastamosing trans- verse ridges; depressions between ridges filled with accreted material, often of contrasting hue, producing distinctive reticulate appearance. Pseudepipleural margin narrowly explanate in anterior fifth, becoming angulate posteriorly; pseudepipleuron flat, granulate and opaque; epi- pleuron subparallel to level of metacoxa, then very abruptly broadened to humerus. Metaster- num finely, sparsely punctate with few very fine, short setae; abdominal sternites 1-3 alutaceous, moderately densely set with very fine, muricate punctures bearing minute, appressed setae. Profemur and mesofemur set with fine, short, appressed setae; protibia with spatulate, distally rounded apical process extending to apex of 1st or 2nd tarsomere (as in Fig. 81); outer margin with row of spinules extending about xh-2h dis- tance to apex, separated by 1-2 spinule widths tarsomere length ratios as follows: 3.5:1.2:1.2 1.2:5.1 (pro-); 6.3:2.0:2.0: 1. 9:5.8 (meso-); 8.7:3.3 2.6:6.1 (meta-). Female.— Apical process of protibia extending to apex of 3rd or 4th tarsomere; coxite length ratio to paraproct length = 0.55. Measurements. — EL 7.8-1 1.6 mm; EW 6.8- 9.8 mm; PL 3.0-4.1 mm; PW 5.6-8.3 mm; BD 4.7-6.7 mm. Lectotype Female (BMNH), hereby desig- nated, from Mexico, Jalisco, Guadalajara; 6 syn- types, same data; 4 syntypes from Colima, Za- potlan. (The latter locality actually refers to the Laguna Zapotlan, near Ciudad Guzman, in Ja- lisco (Selander and Vaurie 1962).) Diagnosis.— Eusattus venosus is similar in most features, including the broad labium, to E. mexicanus Champion. It is distinguished by the anteriorly explanate pseudepipleuron (angulately rounded in mexicanus) and the very abruptly expanded epipleural base (much more gradually broadened in mexicanus). E. venosus differs from E. pons Triplehorn in the posteriorly rounded pseudepipleuron (explanate in pons) and in the reticulate elytral sculpturing (coarsely rugose in pons). Distribution (Fig. 87). — Nayarit and Jalisco, Mexico. Additional Material Examined. — Mexico: Nayarit: 8.7 mi E San Bias, XII-1962 (1); Tepic (1); 24 mi SE Tepic, VII- 3-1962 (1); 26 mi SE Tepic, XI-23-1948 (1); 30 mi SE Tepic. XI-23- 1 948 ( 1 ); Xtlan del Rio, IX-22- 1952(12). Jalisco: mtns. N Ajijic, 7500', IX-20-1964 (1); Acatlan, VIII-28-1941 (2); Guadalajara, summer 1971 (1); 111-20-1903 (1); 15 mi NE Guadalajara, IX- 17- 1970 (2); 3 mi N Guadalajara, X-22-1950 (1); El Molino, VIII- 1 1-1941 (1); La Quemada, X-30-1958 (1): 8.3 mi NE Tala, Puente Tortugas, 5500', XH-27-1971 (1); Volcan Tequila, 10-14 km SSW Tequila, 2143 m, XI-8-1974 (1). Eusattus muricatus Species Group Head muricately punctate or tuberculate; ep- istomum shallowly to deeply and angulately emarginate medially, scarcely emarginate or en- tire at lateral epistomal sutures. Eye very weakly reniform, dorsal lobe distinctly larger than ven- tral. Antennae subfiliform or moniliform basally, segments 6-10 gradually enlarged, scape subgla- brous or with basal 1-2 segments sparsely setose. Mentum 1.4-1.7 x broader than long, anterior angles acute, broadly or very narrowly rounded; anterior margin moderately to deeply emargin- ate; submentum about '/2-% width of mentum, very narrow, sometimes obsolescent; gular su- tures strongly convergent anteriorly, separated by no more than 'A basal width of mentum. Pronotum very strongly convex, broadest at base, lateral margins arcuate, narrowly beaded and moderately explanate, becoming more broadly so before posterior corners; anterior cor- ners rounded, right-angled; disk tuberculate, at least in marginal quarters; hypomeron glabrous, polished, longitudinally furrowed, with dense submarginal fringe of very long, projecting setae, sparse, erect setae along anterior margin and oc- casional setae on coxal cowling; prosternum be- fore coxae much shorter than prosternal process; prosternal process 1.5-1.7 x wider behind than between coxae, flat or weakly convex. Elytra very strongly convex, lateral margins arcuate, broadest near middle or slightly before; pseudepipleural margin broadly rounded, un- defined; epipleuron gradually narrowed from hu- merus to middle of 1st abdominal sternite, then 64 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 • MEXICANUS O VENOSUS ▲ PONS Figure 87. Known distribution of Eusattus mexicanus, E. venosus, and E. pons. subparallel to 5th sternite and narrowed to apex; mesosternum between coxae wider than proster- nal process, deeply excavate; metasternal suture obsolescent, less than Vi length of metasternum, or absent. Profemur and mesofemur bearing dense fringes of long, projecting setae; protibia with row of spinules extending about '/: distance to apex of outer margin, spinules subcontiguous or separated by about 1 spine width basally, be- coming irregular and separated by 2-4 spine widths distally. Metendosternite with arms fused with meta- notum apically, with mesocoxal inflections ba- sally; mesendosternite with arms extending about Vi distance to elytral articulation, flanged basally; spermathecal tubes short, thick (as in Fig. 7); spatulate process of ovipositor nearly straight, rounded apically; aedeagus with apex bluntly rounded; coxite length ratio to paraproct length variable. Eusattus dilatatus LeConte Eusattus dilatatus LeConte, 1851:132; 1858:37; 1866:112: Lacordaire 1859:221; Horn 1870:294; Doyen 1974b:85 (biology); 1977:5 (synonymy). Eusattus muricatus, Horn 1883:304 (in part); Papp 1961:1 16. Sphaeriontis dilatata, Casey 1908:76. Coelosattus fortinen Blaisdell, 1927:167. Eusattus (Sphaeriontis) muricatus, La Rivers 1949:180 (in part). Sphaeriontis dilatatus, Triplehorn 1968:378. Eusattus fortineri. Doyen 1974b:85. Subglobular, black beetles with densely setose venter and arcuate hind tibiae. Male. — Frons and vertex set centrally with very strongly asperate punctures or tubercles about 2 eye facets in diameter, separated by 1- 2 puncture diameters, becoming tuberculo-punc- tate laterally and usually along epistomal suture; epistomum with lateral lobes tuberculo-punc- tate, medial lobe tuberculo-punctate near suture, becoming nearly smooth near anterior margin; deeply, broadly and evenly emarginate medially, barely indented at lateral sutures; epistomal su- ture impressed, sometimes obscured by sculp- turing. Antenna subequal to foretarsus in length; segment length ratios as follows: 2.2: 1.3:1.1 :0.8: 0.7:0.6:0.6:0.6:0.6:0.7:0.9; terminal segment about 1.2 x broader than long, apically rounded or broadly angulate, slightly asymmetrical. Men- turn about 1 .5 x broader than long; anterior bor- DOYEN: SYSTEMATICA OF EL'S.MTS AND COSISA III S 65 der moderately deeply and angulately emargin- ate; lateral borders strongly raised in basal half. Pronotum about 2.2 x broader than long; disk shining, immaculate centrally and anteriorly, with irregular patches of low, rounded tubercles par- amedially near posterior border, becoming reg- ularly set in lateral sixths with oval tubercles 2- 3 eye facet diameters in length, separated by 1 tubercle space or less; tubercles subtended by short, declined setae, especially evident near lat- eral borders. Presternum and prosternal process rugulose to punctato-rugose; presternum set with short, declined setae, often interspersed with few long, erect setae; process moderately densely clothed by long, erect setae, unmargined, round- ed posteriorly. Elytra 1.1—1.2 x longer than broad; disk weak- ly rugulose, set with broad, flattened, often some- what indistinct tubercles basally, these becoming distinct, elongate and asperate posteriorly, about 2-3 eye facet diameters in length, separated by 1 tubercle length or less; epipleuron densely punctate, densely set with long, yellow projecting setae from base to apex. Thoracic venter densely set with long, projecting setae, except in central metasternum. Protibia very broadly flanged (Fig. 88), apical process appearing short, extending about to apex of basal tarsomere; posterior bor- der with fringe of long, projecting, close-set setae; middle and hind tibiae arcuate, with inner sur- face fringed with moderately long, projecting se- tae (Figs. 89-90); hind tibia strongly flattened in cross-section; tarsomere length ratios as follows: 2.9:0.8:0.8:0.7:2.5 (pro-); 5.4:1.6:1.4:1.2:5.5 (meso-); 5.6:1.7:1.5:3.5 (meta-). Female. — Coxite length ratio to paraproct length = 0.38; otherwise differs as stated in ge- neric description. Measurements. — EL 6.9-10.6 mm; EW 6.0- 9.0 mm; PL 2.4-3.7 mm; PW 5.4-8.2 mm; BD 4.7-7.2 mm. Holotype Female, MCZ. The type lacks an- tennae, mouthparts, and terminal abdominal sternites, but is easily recognizable from the re- maining fore and hind left legs. Type Localities.— Of dilatatus, vicinity of the lower Colorado River; fortineri. Grey's Well, near Yuma, in Imperial Co., California. Diagnosis. — Eusattus dilatatus is distin- guished from all other species by the peculiar structure of its legs and the extremely short an- tennae. In other features it is similar to E. muri- catus LeConte. !_-. , , 88 Figures 88-90. Posterior aspect of left legs of Eusattus dilatatus. 88. Prothoracic. 89. Mesothoracic. 90. Metathoracic. Distribution (Fig. 91) and Habitat. — Adults and larvae are entirely restricted to aeolian sand formations, ranging from Riverside Co., Cali- fornia, south to Puerto Penasco, Sonora, Mexico. The beetles are largely subterranean, aggregating in the sand beneath clumps of Larrea or other 66 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 • DILATATUS ■ HIRSUTUS A PHREATOPHILUS Figure 9 1 . Known distribution of Eusattus dilalatus, E. hirsutus, and E. phreatophilus. shrubs. Unlike E. muricatus, which is regularly found on the sand surface in large numbers, di- latatus apparently ventures forth seldom, and is not readily trapped in pitfalls. The distributions of dilatatus and muricatus are almost entirely allopatric (Figs. 9 1 and 99), converging only in the sand hills near Blythe and in the northern Coachella Valley, Riverside Co. Additional Material Examined. — The great majority of specimens have geen collected from the Algodones Sand Dunes in SE California, mostly from the vicinity of Glamis, where a paved road crosses the dunes. These records, spanning the seasons and numbering more than 250 individuals, are not included here. Arizona: Yuma Co.: sand dunes E Yuma, IV- 4-1956 (2); 10-1 5 mi NE Yuma, IV-3/4-1956, sifted from sand under plants (8). California: Imperial Co.: Kane Springs, III- 30-1952 (7); Salton Sea, VIII-7-1948 (1); Superstition Mtn., IX-23-1964 (1); Westmoreland, V-6-1933 (2); Winterhaven, 111-25- 1 956, under creosote (2). Riverside Co.: 3 mi W Blythe, IV-23-1972, pitfall (1); 9 mi N Blythe. IV- 10- 1968 (1); Coa- chella Vy., IV-22- 1955 (1). San Diego Co.: Borego, I V-26- 1954 (5); IV-28- 1955(1); Borrego Vy., I V-29- 1961(1); Borrego State Park, dunes nr. dump, under Larrea, IX-27- 1 980 (2). Mexico: Sonora: Puerto Penasco, XI-24-1955 (6); 1 km W Puerto Pe- nasco, 111-24-78, beach dunes (3); 50 mi SW Sonoyta, III- 12- 1973 (20); 55 mi W Sonoyta, XII-27-1966 (9); 10 mi N Sotelo nr. Bahia Adair, 111-13-1973(5). Eusattus hirsutus, new species Strongly convex, broadly oval, black beetles with long, slender setae sparsely covering the ely- tra. Male. — Frons set with rounded tubercles about 2 eye facets in diameter, separated by 1-2 tu- bercle diameters centrally, becoming almost con- tiguous near eyes and epistomal suture and spars- er on epistomum; intertubercular spaces set with punctures Vy-lh eye facet in diameter, separated by about 1 puncture diameter. Epistomum mod- erately and angulately emarginate medially, scarcely indented at lateral sutures; epistomal su- ture obscured by sculpturing, especially medi- ally. Antenna subequal to forefemur in length; segment length ratios as follows: 3.2: 1 .2: 1 .9: 1 .6: 1.6:1.4:1.3:1.3:1.2:1.2:1.5; terminal segment 1.25 x longer than broad, asymmetrical, with an- gulate apex. Mentum about 1.6 x broader than long; anterior margin moderately emarginate (Fig. 54); lateral margins weakly raised in basal half. Pronotum 2.2-2.3 x broader than long; disk with duplex sculpturing, set centrally with larger, muricate punctures about 2 eye facets in diam- eter, separated by about 1-3 puncture diameters, becoming more strongly muricate and setigerous laterally, until distinctly tuberculate in lateral sixths; smaller punctures about vh eye facet in diameter, separated by 1-2 puncture diameters, visible centrally, obscured laterally; prosternum and prosternal process weakly punctato-rugu- lose, sparsely set with long, slender, erect or de- clined setae; prosternal process margined or not, attenuate posteriorly with truncate or rounded apex. Elytra about 1.3 x longer than broad; disk ru- gulose, set with tubercles about 2-3 eye facets in diameter, separated by 2-3 tubercle diameters anteromedially, becoming closer laterally and on declivity and finer, sparser near epipleural su- ture; tubercles subtended by long, slender, erect, somewhat woolly setae; epipleuron asperately and setigerously punctate nearly to apex. Thoracic sternites sparsely set with long, slender, project- ing setae. Protibia with apical process extend- ing to apex of 2nd tarsomere; posterior border with comb of 7-9 short, stout setae (as in Fig. 96); middle and hind tibiae subcylindrical, evenly spinose; tarsomere length ratios as fol- lows: 4.3: 1.3: 1.3: 1 .2:3.3 (pro-); 6.0:2.2:2.0: 1 .7:4.0 (meso-); 8.3:2.7:2.2:4.7 (meta-). Female. — Differs from male as stated in ge- neric description; ovipositor not examined. Measurements. — EL 7.6-8.6 mm; EW 6.1- 6.5 mm; PL 2.6-2.9 mm; PW 5.9-6.3 mm; BD 4.7-5.1 mm. Holotype Female (CAS) from Nevada, Min- DOYEN: SYSTEMATICS OF EUSATTUS AND COMSAIll'S 67 eral Co., Rhodes Salt Marsh sand dunes, 11-27- 1973 (D. Giuliani). Diagnosis.— Eusattus hirsutus shares charac- ters of E. muricatus LeConte and E. phreato- philus Doyen, differing from both in its setose elytra and pronotum. From muricatus it further differs in having the protibial fringe of setae short and comblike (usually long and slender in muri- catus), the antennae relatively shorter and the ventral epistomal surface subglabrous (setose in muricatus). From phreatophilus it differs in the moderately emarginate mentum (deeply emar- ginate in phreatophilus). Distribution (Fig. 91). — Esmeralda and Min- eral cos., Nevada. Additional Material Examined. — Nevada: Esmeralda Co.: 4 mi W Coaldale, sand dunes. I- 1 3- 1973 (2); 10 mi E Coaldale, sand dunes, 11-19-1973 (1). Eusattus muricatus LeConte (Figure 92) Eusattus muricatus LeConte, 1851:132. Subglobular, black beetles with glabrous or subglabrous, tuberculate elytra. Male. — Frons set with round or elongate tu- bercles about 1-1 Vi eye facets in diameter and separated by 1-3 puncture diameters, becoming closer laterally, usually subcontiguous near eyes and sparser on epistomum. Epistomum narrow- ly, moderately deeply and angulately emarginate medially (Fig. 59), scarcely or not indented at lateral sutures, edges upturned; epistomal suture fine, usually faintly impressed, often obscured by sculpturing. Antenna subequal to forefemur in length; segment length ratios as follows: 3.5:1.6: 1.6:1.4:1.5:1.4:1.4:1.3:1.3:1.2:1.3; terminal seg- ment about 1.2 x longer than broad, asymmet- rical, with angulately rounded apex. Mentum about 1 .4 x broader than long; anterior border moderately deeply, arcuately emarginate (Fig. 54), lateral margins upturned in basal half. Pronotum 2.2-2.4 x broader than long; disk with duplex sculpturing; set centrally with strongly muricate punctures about 1-2 eye facets in di- ameter, separated by 2-3 (1-6) puncture diam- eters, gradually becoming tuberculate laterally; tubercles slightly larger than central punctures, separated by 1-2 diameters, usually setigerous along lateral margin; smaller punctures Vy-lA eye facet in diameter, separated by 1-2 puncture di- ameters, or absent; prosternum and prosternal Figure 92. Eusattus muricatus muricatus LeConte. process punctato-rugulose, sparsely set with moderate to long, slender, inclined setae; pro- sternal process unmargined, at least posteriorly; broadly rounded posteriorly. Elytra 1. 1-1.3 x longer than broad; disk ru- gulose, set with irregular tubercles about 2-4 eye facets in diameter, separated by 1-4 tubercle di- ameters anteromedially, becoming regular and closer on declivity, sparser and usually setigerous near epipleural suture; setae short to moderately long, weakly to moderately inclined; epipleuron asperately and setigerously punctate, more densely so in basal half. Thoracic sternite densely set with long, slender, projecting setae. Protibia with apical process extending to apex of 3rd or 4th tarsomere (Fig. 93); tarsomere length ratios as follows: 2.9:0.8:0.7:0.7:2.5 (pro-); 4.5: 1 .8: 1 .5: 1.3:3.5 (meso-); 5.6:2.2:1.5:3.9 (meta-). Female. — Coxite length ratio to paraproct length = 0.34; otherwise differs from male as stated in generic description. 68 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figures 93-95. Posterior aspect of left legs of EusattUi muricatus muricatus. 93. Prothoracic. 94. Mesothoracic. 95. Metathoracic. Holotype Female (MCZ) from Oregon. Diagnosis. — Eusattus muricatus is distin- guished from E. puberulus LeConte and /:'. hir- sutus Doven bv the absence of setae on the elvtral disk. The mentum of muricatus is much more shallowly emarginate than in E. phreatophilus Doyen. Habitat.— E. muricatus is restricted to very sandy, friable substrates, especially aeolian dunes. In the Mojave Desert and Great Basin, essen- tially every drift of loose sand is inhabited, as well as all the major dune systems. Dispersal along sandy arroyos must account for the broad distribution, but the beetles are very rarely found in such situations. At present most of the pop- ulations appear to be effectively isolated, often by many kilometers of rocky or clay substrate. Adults and larvae are present throughout the year, at least in southerly parts of the range. Adults appear to be less subterranean than those of E. dilatatus, and large numbers are present on the sand surface most nights during the warm season. Even during the cool months, occasional indi- viduals appear on the surface (Andrews et al. 1979). Adults are generalist herbivores and scav- engers (Rogers et al. 1978), feeding mainly on the ground, but also climbing onto low herbs or shrubs to chew the leaves or flowers. Variation. — Within populations, the largest individuals of either sex are about 1 .3 x the size of the smallest. Minor variation in sculpturing and setation are ubiquitous, as in other Eusattus. The setae fringing the posterior foretibiae are occasionally short and bristle-like. Geographic variation is most evident in char- acters reflecting body size, which varies almost by a factor of 2 among localities. Throughout most of the range, body size is inversely corre- lated with altitude and latitude. The largest bee- tles inhabit dune systems situated at relatively low elevations in southern California and Ne- vada. These correspond to E. fulvescens Casey and E. latissima Casey. Body size is relatively large in samples from low-elevation basins in the Death Valley region, and significantly less at higher elevations a short distance away. The most extreme case involves Eureka Valley (915 m), where the beetles are large, and Deep Springs Valley (1530 m), where the beetles are much smaller— these localities are separated by less than 40 km. Relatively large size recurs in samples from a few hundred meters elevation in the Co- lumbia River basin. Beetles from the high-pla- teau country of Nevada, Utah, northeastern Ar- izona, and New Mexico are uniformly small. These correspond to E. acomana Casey. Because DOYEN: SYSTEMAT1CS OF EUSA TTVS AND ( OS ISA III S 69 Figures 96-98. Mctathoracic. Posterior aspect of left tibiae of Eusattus muricatus diablocnsis. 96. Prothoracic. 97. Mesothoracic. 98. of the strong dependence of size on altitude, all of these populations are included in the subspe- cies E. muricatus muricatus. In this subspecies, the foretibia are fringed posteriorly with long, slender setae, and the middle and hind legs usu- ally bear a few slender setae (Figs. 93-95). In the vicinity of San Felipe, in northern Baja California, there occur a series of populations which are disjunct from those in California (Fig. 99). The intervening area, in Riverside, San Die- go, and Imperial cos., is occupied by E. dilatatus. The Baja California populations of muricatus oc- cur at low elevations (10-400 m), but are small in body size. The foretibial setal fringe of these beetles is reduced to a sparse comb of short bris- tles, and the setation on the middle and hind legs is reduced (Figs. 96-98). These Baja California populations are included in the subspecies E. muricatus diablocnsis. Eusattus muricatus muricatus LeConte Eusattus muricatus LeConte, 1851:132; 1866:112; Lacor- daire 1859:221; Horn 1870:294; 1883:305; 1894:423 (key); Cockerell and Fall 1907:202 (distribution); Hatch 1965: 137; Doyen 1972:369 (morphology); 1977:6 (synonymy); Tschinkel and Doyen 1976:331-335 (biology): Rogers et al. 1978 (biology). Sphaenontis muricata, Casey 1908:75. Sphaenontis dilatata, Casey 1908:75. Sphaeriontis acomana Casey, 1908:76. Sphaeriontis latissima Casey, 1924:310. Sphaeriontis fuhescens Casey, 1924:310. Eusattus (Sphaeriontis) muricatus, La Rivers 1949:180 (in part). Eusattus (Sphaeriontis) acomana, La Rivers 1949:180 (syn- onymy). Eusattus (Sphaeriontis) latissima. La Rivers 1949:180 (syn- onymy). Eusattus (Sphaeriontis) fuhescens, La Rivers 1 949: 1 80 (syn- onymy). Sphaeriontis muricata, Triplehorn 1968:378. Sphaeriontis acomanus, Triplehorn 1968:378. Sphaeriontis latissimus. Triplehorn 1968:378. 70 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figure 99. Known distribution of Eusattus muricatus and /•.'. puberulus. Single records of muricatus muricatus from Nogalcs, Santa Cruz Co., Arizona; Long Ridge, Humboldt Co., and Sacramento, Sacramento Co.. California; and Prince of Wales Island. Alaska, are almost certainly in error, and are not plotted. A single record from Montana lists no specific locality. DOYEN: SYSTEMATICS OF EUSATTUS AND CON/SATTUS 71 Body length (elytra plus prothorax) 8.2-16.4 mm; foretibia with posterior fringe of long, slen- der setae (Fig. 93). Measurements. — EL 6.1-12.4 mm; EW 5.2- 9.8 mm; PL 2.1-4.0 mm; PW 4.7-9.1 mm; BD 4.0-8.2 mm. Holotype Female (MCZ) from Oregon. Type Localities.— Of acomana, New Mexico; fulvescens, Owens Lake, Inyo Co., California; la- tissima, Palm Springs, Riverside Co., California. Distribution (Fig. 99).— Eusattus muricatus muricatus ranges discontinuously from east-cen- tral Washington south through the arid, eastern portions of Oregon and California and east to southwestern Colorado and western Texas. Dis- junct populations are known from a few speci- mens from near Colorado Springs and Fort Col- lins, Colorado. Eusattus muricatus diabloensis, new subspecies Body length (elytra plus prothorax) 8.9-11.8 mm; foretibia with posterior row of 7-9 short, stout bristles (Fig. 96). Measurements. — EL 6.7-9.0 mm; EW 5.7- 7.3 mm; PL 2.0-2.8 mm; PW 5.3-6.5 mm; BD 4.3-5.7 mm. Holotype Female (CAS) from Mexico, Baja California Norte, San Felipe, III-5-1963, P. H. Arnaud, Jr. Paratypes from Mexico, Baja Cali- fornia Norte, San Felipe, IV-27-1954, A. Ebel- ing, (1, EME); 45 mi NW San Felipe, 1000', 1-27/ 28-1970, E. L. Sleeper and J. A. Gruwell (3, CSULB); Los Medanos Dunes, 66 mi N San Fe- lipe, 50', 1-16-1976, R. L. Aalbu (1, RLA); km 88, 60 mi S Mexicali, VI-3/4-1961, H. F. How- den (1, CNC); sand dunes at S end Diablo Dry Lake, San Felipe Vy., IV-6-1973, J. Doyen (7, EME). Distribution (Fig. 99). — Extreme northeast- ern Baja California. Eusattus phreatophilus, new species Strongly convex, broadly oval, black beetles with glabrous, finely tuberculate elytra. Male. — Frons set with asperate punctures about 1 V2 eye facets in diameter, separated by 1- 3 puncture diameters centrally and on epi- stomum, disappearing on vertex and becoming denser along epistomal suture and near eyes. Ep- istomum moderately and angulately emarginate medially, scarcely or not indented at lateral su- tures, edges upturned; epistomal suture fine, faintly impressed, often obscured, especially me- dially. Antenna subequal to forefemur in length; segment length ratios as follows: 2.8: 1.1:1.3:1.2: 1.2:1.2:1.0:1.0:1.0:0.9:1.2; terminal segment subtrapezoidal, apically angulate, asymmetrical. Mentum about 1.3 x broader than long, medial half of anterior border deeply emarginate (Fig. 55), lateral margins strongly upturned in basal half. Pronotum 2. 1-2.2 x broader than long; disk with duplex sculpturing; set centrally with larger punctures about as large as eye facets, separated by about 2-6 puncture diameters, becoming muricate laterally until forming distinct tubercles about 1 1/2— 2 eye facets in diameter laterally, sep- arated by about 1-2 diameters and supertending short, fine, appressed setae; smaller punctures shallow, about lk-lh eye facet in diameter, sep- arated by about 1-2 puncture diameters, ob- scured laterally. Presternum and prosternal pro- cess weakly rugulose, the former sparsely set with moderately long, erect setae; process unmar- gined, rounded or truncately rounded. Elytra about 1.3 x longer than broad; disk weakly rugulose, basally set with tubercles or very strongly muricate punctures about 2-5 eye facets in diameter, separated by 2-4 tubercle diameters anteromedially, becoming denser laterally, and denser, more strongly tuberculate and subtended by short, fine setae on declivity; epipleuron se- tose in basal Vi-Vi. Thoracic sternites sparsely set with moderately long, declined setae laterally. Protibia with apical process extending to apex of 1 st or 2nd tarsomere; posterior border with comb of 7-9 short, stout setae (Fig. 100); middle and hind tibiae subcylindrical, nearly straight, evenly spinose (Figs. 101-102); tarsomere length ratios as follows: 3.2:0.8:0.7:0.7:2.9 (pro-); 4.0: 1 .9: 1.5: 1.3:3.5 (meso-); 5.5:2.2:1.7:3.7 (meta-). Female.— Coxite length ratio to paraproct length s 0.50; otherwise differs as stated in ge- neric description. Measurements. — EL 5.9-9.9 mm; EW 4.6- 7.5 mm; PL 2.2-3.3 mm; PW 4.6-7.3 mm; BD 3.7-6.1 mm. Holotype Female (CAS) and 3 paratypes (EME) from California, San Bernardino County, 9 mi S Baker, sand dunes S of Zzyzx Springs, IV- 22-1977, M. E. Buegler. Additional paratypes (EME, RLA) as follows: same data, IV-25-1977, in sand under Cleomella, J. Doyen (16); same data, IV-27-1977, J. Doyen (J. D. lot 77D2) (25); 72 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figures 100- thoracic. 102. Posterior aspect of left tibiae of Eusattus phreatophilus. 100. Prothoracic. 101. Mesothoracic. 102. Meta- same data, IV-28-1977, J. Doyen, ex pitfalls (4); same data, IV-22/24-1977, R. L. Aalbu (9). Diagnosis. — Eusattus phreatophilus is most similar to E. hirsutus Doyen, differing as de- scribed under the latter. It is similar to E. muri- catus LeConte, differing in the very deeply emar- ginate mentum (moderately emarginate in muricatus; see Figs. 54-55). Distribution (Fig. 91) and Habitat. — Adults and larvae occur in sandy substrates, usually about the edges of dry lakes or saline springs, where they hide in the loose sand and debris which accumulate beneath low-growing vegeta- tion. The known distribution, from Nye Co., Ne- vada, south to San Bernardino Co., California, is restricted to undrained basins entirely within the range of E. muricatus. The latter species mainly occupies aeolian dunes, where phreato- philus does not occur. Both species have been collected from intermediate situations such as the dunelets south of Baker Dry Lake, San Ber- nardino Co., and in Saline Valley, Inyo Co., Cal- ifornia. Additional Material Examined. — California: Inyo Co.: Death Valley Natl. Mon., Furnace Creek Camp, 111-30-1949 ( 1 ); Saline Valley: IV-2/3- 1 960 (2), V-7- 1 960 (4), VII- 1 976 ( 1 ); SE end, dead edge of dry lake, 111-30-1976 (22); 1 100', 11-7/ IV-25-1978 (6); salt marsh, 1060', 111-23-1976 (2), IV-1-1976 DOYEN: SYSTEMATICS OF EUSATTUS AND COXISATTL'S 73 (3), V-6-1976 (2), VII-1-1976 (1); artesian spring. IV-24-1975 (1); upper warm spring, 1900', IV-6-1976 (1); dunes, 1000', IV-27- 1974(1); Morning Sun Mine, IX- 1 977 (1); San Bernar- dino Co.: Mesquite Valley, 11-18-1978 (4); Saratoga Springs, III- 1 5- 1 970 ( 1 ), 111-24- 1 980 (2), IV- 1 7- 1 974 (2). Nevada: Clark Co.: Las Vegas, IX- 1975 (1); Moapa floodplain, 1 mi N Glen- dale, VI-1 1-1975 (2); Nye Co.: Ash Meadow, VI-5-1954 (1 1); 15 mi S Lathrop Wells, 11-28-1976 (1). Eusattus puberulus LeConte Eusattus puberulus LeConte, 1854:84; 1858:37; 1866:112; Lacordaire 1859:221; Horn 1870:294; 1883:204; 1894: 423 (key); Papp 1961:116. Sphaeriontis puberula, Casey 1908:77. Eusattus (Sphaeriontis) puberulus. La Rivers 1949:180. Sphaeriontis puberulus, Triplehorn 1968:378. Subglobular, black beetles with pubescent ely- tra. Male. — Frons and vertex smooth centrally, becoming tuberculate laterally and near epistom- al suture; tubercles 1-2 eye facets in diameter, separated by about 1-3 tubercle diameters, den- sest laterally, usually obscured on epistomum, especially medial lobe, sometimes supertending short, flattened setae on frons. Epistomum mod- erately, narrowly and angulately emarginate me- dially, with edges upturned, especially laterally; epistomal suture obscured by cuticular sculptur- ing, especially laterally. Antenna subequal to forefemur in length; segment length ratios as fol- lows: 1.9:0.9:1.0:0.7:0.8:0.8:0.8:0.8:0.9:0.9:0.9; terminal segment about as long as broad, apically rounded, slightly asymmetrical. Mentum about 1.6 x broader than long, moderately and evenly emarginate in middle half of anterior border; lat- eral margins raised in basal third. Pronotum about 2.3 x broader than long; disk set with exceedingly fine punctures separated by about 2 puncture diameters; in lateral sixths set with tubercles 1-2 eye facets in diameter and supertending short to moderately long, flattened setae directed posteromedially; lateral carina mi- nutely fimbriate; presternum and prosternal pro- cess punctato-rugulose, sparsely set with mod- erately long, declined setae; prosternal process margined, evenly rounded posteriorly. Elytra about 1.2 x longer than broad; disk set anteriorly and medially with broad, shallow, as- perate punctures, each bearing a short, flattened seta; gradually becoming more strongly asperate posteriorly and laterally, until distinctly tuber- culate on declivity; epipleuron asperately and se- tigerously punctate in anterior half. Thoracic sternites sparsely set with long setae laterally; Figure 103. Posterior aspect of left foretibia of Eusattus puberulus. abdominal sternites 1-3 sparsely set with very short, fine appressed setae. Protibia with apical process extending to apex of 2nd to 4th tarso- meres; posterior margin with fringe of long, pro- jecting setae (Fig. 103); metatibia subcylindrical, nearly straight, evenly spinose. Tarsomere length ratios as follows: 2.8:0.6:0.6:0.5:2.0 (pro-); 4.3: 1.5:1.3:1.0:2.7 (meso-); 5.2:2.0:1.3:3.3 (meta-). Female. — Coxite length ratio to paraproct length = 0.32; otherwise differs from male as stated in generic description. Measurements. — EL 5.1-6.8 mm; EW 4.2- 5.5 mm; PL 1.7-2.2 mm; PW 3.9-5.1 mm; BD 3.5-4.1 mm. Holotype Female (MCZ) from Texas, be- tween Laredo and Ringgold Barracks. Diagnosis.— Eusattus puberulus is most sim- ilar to E. muricatus LeConte, differing in its an- teriorly punctate, setose elytra (tuberculate, gla- brous or nearly so in muricatus). E. hirsutus Doyen has setose elytra, but the setae are much longer and the elytra tuberculate. E. minimus Doyen is superficially very similar, but belongs to the convexus species group and differs from puberulus in many details. Distribution (Fig. 99) and Habitat. — Re- stricted to the sand dunes and sand flats in south- eastern Texas from the vicinity of Laredo north to Victoria and east almost to the Gulf, but not on dunes immediately adjacent to the coast. Nearly all the interior sand-dune habitat in southeastern Texas is enclosed in large ranches with very restricted access. Consequently, E. puberulus remains very poorly represented in collections. Additional Material Examined.— Texas: Brooks Co.: La- guna Salado, X-27 (1); Goliad Co.: Goliad (2); Jackson Co.: X-25-1975 (1); Kennedy Co.: Riskin Ranch, El Paistle, IV-22- 1975/XI-20-1979 (5); Kleberg Co.: La Paloma Ranch, V-19- 74 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figures 104-106. Posterior aspect of left legs of Eusattus arenarius. 104. Prothoracic. 105. Mesothoracic. 106. Meta- thoracic. 1975 (1); San Patricio Co.: Welder Wildlife Refuge, IX- 13- 1979(1); Webb Co.: 15 mi SE Laredo, IX-20/X- 17- 1980(1 1); 19 mi SE Laredo, X- 17- 1980 (1). Eusattus ciliatus Species Group Very strongly convex or globular beetles with finely tuberculate elytra. Head and pronotum with simple or duplex punctation; epistomum mod- erately to deeply and broadly emarginate me- dially; slightly to deeply indented at lateral su- tures, margin weakly to moderately reflexed; epistomal suture distinct, weakly impressed; eye ovoid or very weakly reniform, scarcely or not indented by epistomal canthus; antenna gradu- ally enlarged to apex, or subfiliform to segment 7 or 8, last 3 or 4 segments enlarged as a weak club; terminal segment symmetrical, ovoid or attenuate. Scape bearing several setae at least as long as 3rd segment; flagellum distinctly pubes- cent (Fig. 34); mentum subtrapezoidal, flat or weakly convex, unsculptured except for few punctures bearing long slender erect setae, an- terior angles slightly acute, narrowly rounded; submentum slightly broader than base of men- tum, gula narrowed anteriorly, V6-V3 width of mentum. Pronotum strongly convex, streamline with contour of elytra, lateral margins narrowly to broadly explanate; hypomeron glabrous, pol- ished except for moderate to dense submarginal fringe of very long, projecting setae and sparse, long setae along anterior margin; sternum opaque, set with long, erect setae; antecoxal length greater than length of prosternal process; prosternal pro- cess about Vi as wide between as behind coxae, rounded posteriorly, margined, at least obscurely so, sparsely set with long, erect setae. Elytra with epipleuron gradually narrowing from base to apex; pseudepipleuron rounded, un- defined; mesosternum between coxae at least as broad as prosternal process, scarcely to moder- ately excavate; metasternal suture V2— 7/s length of metasternum; metasternal sculpturing variable; intercoxal process narrow, acutely to obtusely rounded (Fig. 42). Profemur and mesofemur bearing dorsal fringe of long, projecting setae, shorter setae along ven- tral surface; protibia with attenuate, narrowly rounded apical process extending to apex of 1st to 3rd tarsomere. Metendosternite with arms fused with meso- notum, not fused with mesocoxal inflections; mesendosternite with arms extending about % distance to mesepisternal process, enlarged ba- sally as muscle attachment flanges; spermathecal tubes 2 or 4 long, slender (Fig. 12), unbranched DOYEN: SYSTEMATICS OF EUSATTUS AND CONISATTUS 75 or paucibranched; aedeagus with bluntly round- ed apex; ovipositor setose, paraproct not ex- panded beneath coxite; spatulate processes vari- able; coxite length ratio to paraproct length variable. Eusattus arenarius, new species Strongly convex, subglobular, dark brown to black beetles with sparsely, very finely tubercu- late elytra. Male.— Similar to E. ciliatus, except in the following characters: Head and pronotum simply sculptured, or with smaller punctures barely vis- ible at 50 x ; frons asperately punctate or tuber- culate posteromedially, tuberculate laterally near eyes, becoming punctate toward epistomal su- ture and on epistomum; tubercles/punctures about '/2 eye facet in diameter medially, separated by 2-4 puncture diameters, coarser and closer laterally and near epistomal suture; epistomum weakly to moderately indented at lateral sutures; epistomal canthus usually contiguous or nearly so with anterior margin of eye; antenna gradually enlarged to apex; segment length ratios as fol- lows: 2.4:1.2:1.7:1.5:1.4:1.4:1.2:1.2:1.2:1.1:1.4; mentum with lateral borders arcuate. Metaster- nal suture as long as metasternum or nearly so; intercoxal process nearly right-angled or obtuse; abdominal sternites 1-3 exceedingly finely punc- tate medially, with few, short setae laterally; metafemur subglabrous or with few, short setae; mesotibia and metatibia with short, sharp spines, without long, slender setae (Figs. 104-106); dor- sal surface of basitarsus of middle and hind legs subglabrous or sparsely set with short, appressed setae; aedeagus similar to that of ciliatus. Female. — Spermathecal tubes 2; ovipositor similar to that of E. ciliatus, but spatulate pro- cesses not upturned; 7th tergite weakly emargin- ate. Measurements. — EL 7.1-9.2 mm; EW 5.5- 7.0 mm; PL 2.3-2.9 mm; PW 5.1-6.5 mm; BD 4.5-6.0 mm. Holotype Male (CAS) and 2 paratypes from Mexico, Baja California Norte, Miller's Landing, 111-29- 1973, sea level, J. Doyen and S. L. Szerlip; additional paratypes as follows: Millers Landing, IV-6-1976, J. Doyen, P. Rude, R. Morrison (14, EME); 1.8 km SE Miller's Landing, beachdune, V-27/28-1973, E. L. Sleeper (9, CSULB); 2 mi S Ejido Morelos, VII-5-1977, sand dunes, J. K. Aalbu (4, RLA); 1 1 .3 km N Guerrero Negro, VII- 1977, sand dunes, J. K. Aalbu (13, EME, RLA); 9 km N Guerrero Negro, IX-8- 1 977, sand dunes, E. Fisher, R. Westcott (8, EME, CSULB); 6 mi N Guerrero Negro, VIM- 1979, A. Hardy, F. Andrews, D. Giuliani (164, EME, CDFA); 6 mi N Guerrero Negro, III- 16- 1981, F. Andrews, D. Faulkner (9, CDFA); 7 km N Guerrero Negro, 111-27-1975, sand dunes, R. L. Aalbu (4, RLA); 5 km N Guerrero Negro, VIII-25-1975, sand dunes, R. L. Aalbu (2, RLA); Baja California Sur, 7 mi W Guerrero Negro, IV-7-1976, J. Doyen, P. Rude, R. Morrison (4, EME); 7 mi SE Guer- rero Negro, IV-8-1976, J. Doyen, P. Rude, R. Morrison (3, EME); 1 km S Guerrero Negro, X- 18- 1977, 30 m, D. E. and W. R. Breedlove (7, CAS). Diagnosis.— Eusattus arenarius is similar to E. ciliatus Horn and E. ciliatoides Doyen, dif- fering from both in lacking the long, slender setae on the metafemur, mesotibia, and metatibia. In arenarius, the metasternal suture extends the en- tire length of the metasternum; in the other two, the suture is no more than Vi as long as the meta- sternum. The basal segments of the middle and hind tarsus of arenarius bear only sparse, short, appressed setae; in ciliatus, the setae are more numerous and longer (Figs. 1 12 and 1 13). Distribution (Fig. 107) and Habitat. — Like most other members of the ciliatus species group, E. arenarius is restricted to aeolian sand dunes. A few individuals have been collected as far as 5 km from the coastline; the great majority have come from dunes no more than a few hundred meters inland. All records are from the dune complex around Scammons Lagoon in Baja Cal- ifornia, at the pacific edge of the Vizcaino Desert. Eusattus ciliatoides, new species Strongly convex, subglobular, dark brown to black beetles with sparsely, finely tuberculate ely- tra. Male. — Similar to E. ciliatus, except in fol- lowing characters: Frons with large punctures as large or slightly larger than eye facets, separated by about 1-3 puncture diameters; epistomum slightly to moderately indented at lateral epi- stomal sutures, strongly so in 1 specimen; epi- stomal canthus usually contiguous or nearly so with anterior margin of eye; antenna with seg- ments 9-1 1 forming weak club; segment length ratios as follows: 2.3:1.1:1.7:1.4:1.5:1.4:1.4:1.4: 1.4:1.4:1.7; mentum about 1.5 x broader than 76 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 Figure 107. Known distribution of Eusat tits arenarms. E. ciliatoides, and E. ciliatits in northern Baja California. long, lateral margins arcuate. Pronotum 2.2-2.8 x broader than long; elytra 1.22—1 .33 x longer than broad; posterior margin of anterior tibia with 5- 8 spines; dorsal surface of basitarsus of middle and hind legs sparsely set with short, appressed setae (as in Figs. 105-106). Aedeagus as in Fig. 108. Female. — Spermathecal tubes 2; ovipositor as in Fig. 109; 7th tergite with rounded, medial emargination (Fig. 1 10). Measurements. — EL 5.7-8.4 mm; EW 4.7- 6.5 mm; PL 1.6-2.7 mm; PW 3.9-6.2 mm; BD 4.2-5.4 mm. Holotype Female (CAS) and 3 paratypes (EME) from Mexico, Baja California Norte, dunes 1 mi S Cantamar, 111-20-1972, J. Doyen (J. D. Lot 72C5); additional paratypes (EME) as fol- lows: same locality, XI-28-1975, P. Rude, J. D. Lot 75K1 (1); XII-29-1975, P. Rude, J. D. Lot 75L1 (4); XII-28-1976, P. Rude, J. D. Lot 75L15.1 (9). Diagnosis. — Eusattus ciliatoides is distin- guished from E. ciliatus Horn and E. arenarius Doyen as described in the diagnoses for those species. Distribution (Fig. 107) and Habitat. — Sand dunes and very sandy substrates from Cantamar, below Tijuana, south to the San Quintin dunes. Adults and larvae burrow in the sand, usually beneath the canopies of perennial plants. Both maritime and inland sands are inhabited. In the northern part of its range, E. ciliatoides averages almost as large as E. ciliatus. In the region around Bahia San Quintin, where it is sympatric with ciliatus, ciliatoides is significantly smaller, suggesting character displacement. Additional Material Examined. — Mexico: Baja Califor- nia Norte: Enscnada. VIII-30- 1952 (1); vie. Arroyo Santo To- mas XI-30-1957 (6); Santo Tomas Canyon, X-8-1950 (1); 10 mi S San Vicente, 111-25-1973 (1); Santo Domingo. VIII- 13- 1 954 ( 1 ); 2 mi E Rancho San Salvador, VI-2 1 - 1 973 ( 1 ); Bahia San Ramon, sand dunes, Vl-24- 1977 (2); 8 mi NW San Quin- DOYEN: SYSTEMATICS OF EL'SATTLS AND COMSAT! IS 77 ' 110 09 ",''."',', iV'M Figures 108-1 10. Diagnostic structures of Eusattus ciliatoides. 108. Tegmen (left) and median lobe (right). 109. Ovipositor, ventral (right) and lateral (left). 1 10. 7th abdominal tergite of female. tin, VI-14-1973 (1); 10 mi E San Quintin, IX-4-1955 (1); San Quintin (1); 1 mi NE El Socorro (4); 1 mi N El Socorro (1); El Socorro, 111-30-1961 (1), VI-2 1-1973 (1). Eusattus ciliatus Horn Eusattus ciliatus Horn, 1894:349, 422, 423. Sphaeriontis ciliata, Casey 1908:76; Blaisdell 1943:194. Eusattus (Sphaeriontis) ciliatus. La Rivers 1949:180. Sphaeriontis ciliatus. Triplehorn 1968:378. Convex, oval, dark brown to black beetles with sparsely, finely tuberculate elytra. Male. — Head and pronotum with duplex punctation; frons set with large punctures slightly smaller than eye facets, separated by 1-4 punc- ture diameters, becoming denser along epistomal suture; small punctures about V4-V6 that size, sep- arated by 1-2 puncture diameters; epistomum strongly indented at lateral sutures (Fig. 60); epi- stomal canthus distinctly exceeding eye laterally and separated from it posteriorly by groove; an- tenna gradually enlarged to apex; segment length ratios as follows: 2.6:1.4:2.0:1.9:1.6:1.4:1.4:1.4: 1 .4: 1 .4: 1 .6; terminal segment about 1 .5 x longer than broad, rounded apically. Mentum about 1 .4 x broader than long, weakly arcuate, reflexed in posterior >/2— 2A; anterior border almost straight, with weak medial indentation; gula narrowed to 'A-'A. width of submentum. Pronotum 2-2.3 x broader than long, base strongly bisinuate; lateral edges strongly expla- nate, horizontal; disk set with coarser punctures about 'A eye facet diameter, separated by about 2-6 puncture diameters, becoming coarser and denser laterally and containing moderately long, declined, backwardly directed setae; finer punc- tures V4-V3 size of coarser, separated by about 1- 4 puncture diameters; sternum rugulose; proster- nal process granulose. Elytra 1.3 1-1 .45 x longer than broad, set with posteriorly declined tubercles about as large as eye facets, separated by 2-6 tubercle diameters, becoming denser and setigerous laterally and coarser on declivity; setae longest along epipleu- ron; epipleuron densely set with muricate, setig- erous punctures in anterior half; setae long, di- rected laterad; mesosternal width between coxae subequal to prosternal process width, shallowly excavate; metasternum moderately densely, shallowly and setigerously punctate; setae long, erect; metasternal suture about Vi length of meta- sternum; intercoxal process acutely rounded. Protibia (Fig. Ill) with spinules extending en- tire length of outer margin, separated by about 1 spinule width basally, becoming contiguous apically; inner margin set with about 8 erect spines, alternated with about 4-5 long, slender setae; anterior face set with sparse, submarginal row of long, laterally directed setae; posterior face glabrous, spine field restricted to basal fourth; mesofemur densely set with long, erect setae ex- cept on posterior surface; tibia with posterior surface sparsely set with moderately long setae (Fig. 1 1 2); metafemur with sparse dorsal and ventral fringes of long setae; tibial setation sim- ilar to that of middle legs; dorsal surface of basi- tarsus of middle and hind legs moderately dense- ly set with long, inclined setae (Fig. 113) tarsomere length ratios as follows: 3.8:1.1:1.0 0.8:3.0 (pro-); 5. 1 :2.2: 1 .9: 1 .5:3.8 (meso-); 6.6:2.3 1.7:4.6 (meta-). Aedeagus as in Fig. 1 14. Female. — Spermathecal tubes 2; coxite length ratio to paraproct length s 0.37; ovipositor as 78 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figures 1 1 1-1 13. Posterior aspect of left legs of Eusattus ciliatus. 111. Prothoracic. 112. Mesothoracic. 113. Metatho- racic. in Fig. 1 15; tergite 7 entire or weakly indented medially (Fig. 1 16). Measurements. — EL 7.5-9.5 mm; EW 5.4- 7.0 mm; PL 2.4-2.7 mm; PW 4.9-6.7 mm; BD 4.5-5.7 mm. Holotype Female (MCZ) from Mexico, Baja California Norte, Big Canyon, Tantilles Mtns. Diagnosis.— Eusattus ciliatus is very similar to E. ciliatoides Doyen, but is larger (see Mea- surements), less convex, and more elongate (0.43 < PL/PW < 0.49 in ciliatus; 0.35 < PL/ PW < 0.44 in ciliatoides). In ciliatus, the basi- tarsi of the middle and hind legs are pubescent (Figs. 112-113); in ciliatoides, they are subgla- brous or bear a few short setae (as in Figs. 105- 106). There are additional, subtle differences in configuration of the ovipositor and aedeagus. In ciliatus, the spatulate processes are straight and relatively more slender and attenuate (Fig. 1 1 5), and the aedeagus is more attenuate apically (Figs. 108 and 114). In ciliatoides, the spatulate pro- cesses are noticeably upcurved and slightly broader (Fig. 109). Distribution (Fig. 107) and Habitat.— Known only from the large dune system centered about Bahia San Quintin. The type locality spec- ified by Horn is inland and considerably north, and probably erroneous. Adults and larvae burrow in the sand beneath canopies of plants such as Haplopappus venetus. All individuals have been collected in strongly maritime situations, ranging from low foredunes a few meters from the line of highest tides to aeolian sands a few hundred meters inland. Additional Material Examined. — Mexico: Baja Califor- nia Norte: Colonia Guerrero, XII-23-1974 (1); Santa Maria Beach at San Quintin Bay, XI-2-1958 (3), III- 19- 1972 (16), 111-26-1972 (25), VI- 13- 1973 (12); 4 mi S San Quintin, IX-4- 1976 (2); Santa Maria, N Lagoon, 30°25', VIII-24-1953 (1); 1 mi N El Socorro, VII-14-1979 (20); 5.7 km S Rancho El So- corro, 20 m, 111-20-1975(2). Eusattus dubius LeConte (Figure 1 17) Eusattus dubius LeConte, 1 85 1 : 1 32. Convex, oval, dark brown to black beetles with finely, sparsely tuberculate elytra. Male. — Head and pronotum with duplex or apparently simple punctation; frons with coarser punctures 1-2 eye facets in diameter, separated by 1-3 puncture diameters, usually becoming closer and coarser on the epistomum and dis- appearing toward the vertex; finer punctures as large as XU eye facet in diameter, or barely visible at 50 x , separated by about 1 puncture diameter; epistomum shallowly to deeply emarginate me- dially, slightly to moderately indented at lateral sutures; lateral lobes of epistomal canthus dis- tinctly exceeding and contiguous with eyes; an- DOYEN: SYSTEMATICS OF EUSATTUS AND CON1SATTUS 79 I l'l'\'J Figures 1 14-1 16. Diagnostic structures of Eusattus aliatus. 1 14. Tegmen (left) and median lobe (right). 1 1 5. Ovipositor, ventral (left) and lateral (right). 1 16. 7th abdominal tergite of female. tenna gradually broadened to 10th segment; mentum about 1.5 x broader than long, flat; lat- eral margins nearly straight in anterior half, then convergent and moderately reflexed to base; an- terior border moderately, evenly emarginate; gula narrowed to V4-V3 width of submentum. Pronotum 1.9-2.3 x broader than long, base moderately bisinuate, lateral edges moderately explanate, horizontal, briefly upturned; disk set with coarser punctures about 1-1 Vi eye facets in diameter, separated by 1-4 puncture diameters centrally, sometimes with impunctate areas, be- coming more coarsely and closely punctate lat- erally, where punctures contain short to mod- erately long, declined, backwardly directed setae; finer punctures Va-V& diameter of coarser ones, sometimes invisible at 50 x; sternum and pro- sternal process granulose. Elytra 1.3-1.5 x broader than long; finely, moderately densely tuberculate to punctate; tu- bercles/punctures separated by 2-6 diameters, irregularly set laterally and posteriorly with very short to moderately long setae; epipleuron mod- erately densely set with long, erect setae, densest and longest anteriorly, becoming very sparse in posterior fourth; mesosternal width between cox- ae subequal to prosternal process width, shal- lowly to moderately excavate, densely setose. Protibia (Figs. 118-119) with spines extending %-% distance to apex along outer margin, sep- arated by 1 spine width basally, 1-2 spine widths apically; inner margin set with irregular row of short spines; anterior face sparsely set with short, decumbent setae; posterior face with spine field irregularly occupying basal half along posterior margin; metafemur sparsely set with moderately long declined setae, or subglabrous; tarsomere length ratios as follows: 2.5:0.8:0.7:0.6:2.3 (pro-); 3.0:1.2:1.1:0.8:2.4 (meso-); 4.2:1.6:1.2:2.6 (meta-). Figure 1 1 7. Eusattus dubius dubius LeConte. 80 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figures 118-119. Posterior aspect ofleft forelegs. 1 18. Eusattus dubius dubius. 119. E. d. setosus. Female. — Spermathecal tubes 2; coxite length ratio to paraproct length = 0.30. Holotype Female (MCZ) from vicinity of Colorado and Gila rivers. Diagnosis.— Eusattus dubius is superficially similar to Conisattus rectus Casey, but is easily separated by the characters in the key to genera. The most similar Eusattus in external characters are E. product us LeConte and E. ciliatus Horn. In productus, the apical antennal segment is asymmetrical and angulate (symmetrical, usually rounded in dubius). In both productus and cil- iatus, the subcontiguous spines along the exterior protibial margin extend to the apex (separated by 1-2 spine widths, extending 3/4-% to apex in dubius). Variation.— The populations included in E. dubius vary in size, elytral sculpturing, and de- gree of emargination of the epistomum. This variation is recognized here at the subspecific level. Key to the Subspecies of Eusattus dubius 1. 2(1). Elytra tuberculate or with very strongly muricate punctures 2 Elytra punctate or punctato-rugulose dubius arizonensis, new subspecies Epistomum deeply emarginate (as in Fig. 59) 3 Epistomum very shallowly emarginate (as in Fig. 61) dubius abditus, new subspecies 3(2). Terminal antennal segment 1.2-1.3 x as long as broad, usually rounded api- cally; 3rd segment 1.4-1.6x as long as 2nd segment; dubius dubius LeConte Terminal antennal segment 1.6-2.0 x as long as broad, attenuate apically; 3rd segment 1.6-2.2x as long as 2nd seg- ment dubius setosus, new subspecies Eusattus dubius dubius LeConte Eusattus dubius LeConte, 1851:132; Lacordaire 1859:221; Horn 1870:294; 1883:205; 1894:423 (key); Champion 1884: 75; Casey 1908:66; Doyen 1972:369 (morphology); 1977: 5 (synonymy). Conipinus dubius, LeConte 1862:223; Casey 1908:162; 1924: 311. Eusattus nanus Casey, 1895:613; 1908:66. Conipinus nanus, Casey 1908:162; 1924:31 1. Eusattus oblongulus Casey, 1908:67; 1924:311. Conipinus oblongulus, Casey 1908:162. Conipinus spaldingi Casey, 1 924:3 1 3. Eusattus (Conipinus) nanus, Leng and Mutchler 1927:35. Eusattus (Conipinus) dubius, Leng and Mutchler 1927:35. Eusattus (Conipinus) oblongidus, Leng and Mutchler 1927: 35. Eusattus (Conipinus) spaldingi, Leng and Mutchler 1927: 35. Epistomum deeply emarginate medially, weakly to moderately indented at lateral sutures; antennal segment length ratios as follows: 2.6: 1.2:1.6:1.4:1.4:1.2:1.2:1.1:1.1:1.1:1.2; terminal segment rounded apically or occasionally atten- uate, 1.2-1.5 x longer than broad; elytra tuber- culate or occasionally set with very strongly mu- ricate punctures, sparsely set with short setae becoming longer, denser near epipleural margin. Measurements. — EL 4.4-6.9 mm; EW 3.2- 5.1 mm; PL 1.5-2.2 mm; PW 3.1-4.8 mm; BD 2.6-4.1 mm. Holotype Female (MCZ) from vicinity of Colorado and Gila rivers. Type Localities. — Of nanus, Kern Co., Cal- DOYEN: SYSTEMATICS OF ELSATTLS AND COMSATTUS 81 lfornia; oblongidas, Lancaster, California; spal- dingi, Las Vegas, Nevada. Diagnosis.— Eusattus dubius dubius differs from E. d. arizonensis in its tuberculate elytra (punctate in arizonensis) and less elongate ter- minal antennal segment. Distribution (Fig. 120) and Habitat.— Cen- tral Nevada south and east to southern Utah, south and west through California east of the Sierra Nevada and Peninsular ranges to San Die- go and Imperial cos. Many desert environments are occupied, varying from sandy washes and flats near sea level to stony areas as high as 2000 m. E.d. dubius occurs east of the Colorado River only near Yuma, Arizona. Although unrecorded in Mexico, it undoubtedly occurs in northeastern Baja California. Eusattus dubius arizonensis, new subspecies Epistomum moderately to deeply emarginate medially, weakly to moderately indented at lat- eral sutures; antennal segment length ratios ap- proximately as follows: 2.4:1.0:1.7:1.6:1.4:1.3: 1 . 3: 1 . 1 : 1 . 1 : 1 .0: 1 .4; terminal segment with atten- uate apex, 1 .6-1 .7 x longer than broad; elytra set with shallow, often somewhat obscured punc- tures 1-2 eye facets in diameter, often weakly rugulose, especially on declivity; very sparsely, finely setose, usually appearing subglabrous. Measurements. — EL 4.8-7.6 mm; EW 3.7- 5.7 mm; PL 1.7-2.7 mm; PW 3.4-5.1 mm; BD 2.8-4.4 mm. Holotype Male (CAS) and 8 paratypes from Arizona, Yuma County, Kofa Mtns., Palm Can- yon, 111-28-1961 (C. A. Toschi). Diagnosis.— Eusattus d. arizonensis is distin- guished from the other subspecies by its punctate elytra. Relative length of antennal segments is allometrically related to size. In the smallest in- dividuals of E. d. arizonensis, the 3rd segment is about 1.5 x longer than the 2nd; in the largest specimens, the ratio is 1.8-1.9. Distribution (Fig. 1 20) and Habitat. — Most specimens have been collected east of the Col- orado River, from Maricopa Co. west and north to Yuma and Mojave cos., Arizona. Collection sites north or west of the Colorado River are in southern Nye Co., Nevada, in the Providence Mtns., San Bernardino Co., California, and in eastern Imperial Co., California. At the Nye Co. locality on the Nevada Atomic Test site, speci- mens with sculpturing typical of both d. dubius Figure 1 20. Known distribution of Eusattus dubius. and d. arizonensis have been collected. Habitats range from sandy substrates near sea level to stony areas as high as 1260 m. Additional Material Examined.— Arizona: Maricopa Co.: Gila Bend, IV- 1966 (1); Phoenix, 11-22-1926 (1); South Moun- tain Park, nr. Phoenix IV-25-1 965 (2); Wickenberg, I V-4- 1 964 (1); Pima Co.: Organ Pipe Cactus Natl. Mon., IV-II-1965 (2). 111-24- 1964 (1), V-II-1969 (1); 111-23-1953 (1); Tucson, IV- 17-1968 (1); Yuma Co.: Ehrenberg, 11-12-1939 (2), 11-15-1940 (4); Martinez Lake, IV-29-1961 (3); Palm Canyon. Kofa Mtns.. IV-25-1 960 (2); Quartzite, 11-22-1958 (1). California: Imperial Co.: 0.3 mi S Palo Verde, IV-9-1963 (1); Picacho Peak, IV- 10-1965 (1); San Bernardino Co.: Providence Mtns., Bonanza King Mine, 4100', IV-8-1966 (4). Nevada: Nye Co.: 23 mi W Indian Springs, 1 9 mi SE Lathrop Wells. 111-24- 1970(1): Rock Valley, Nevada Test Site V-27/VI-28-1971 (4). 82 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figure 121. Eusattus pallidus pallidus Doyen. Eusattus dubius abditus, new subspecies Eusattus dubius, Blaisdell 1943:192 (in part). Epistomum barely emarginate medially (as in Fig. 61), scarcely or not indented at lateral su- tures; antennal segment length ratios approxi- mately as follows: 2.4: 1 .4:2.6:2.0: 1 .6: 1 .7: 1 .6: 1 .5: 1 .4: 1 .4: 1 .6; terminal segment rounded or slightly attenuate, about 1.6 x longer than broad; elytra set with strongly muricate punctures about as large as eye facets basally, becoming tuberculate on declivity, very sparsely, finely setose, ap- pearing subglabrous. Measurements. — EL 6.8-8.4 mm; EW 5.0- 5.9 mm; PL 2.2-2.7 mm; PW 4.6-5.6 mm; BD 3.8-4.6 mm. Holotype Male (CAS) and 2 paratypes (EM E) from Mexico, Baja California Norte, Millers Landing, 111-29- 1973, J. Doyen and S. L. Szerlip; 4 paratypes (EME), same locality, IV-6-1971, J. Doyen, P. Rude, R. Morrison. Diagnosis. — Eusattus dubius abditus is distin- guished by its very shallowly emarginate episto- mum. The average body size is larger than in d. dubius or d. arizonensis. Distribution (Fig. 120) and Habitat.— Oc- curs only in the Vizcaino Desert and the region just to the north. The type series was collected on aeolian coastal dunes, where the beetles were active at night. Other available ecological infor- mation also lists sand dunes as the habitat. Additional Material Examined. — Mexico: Baja Califor- nia Norte: 2 mi S Ejido Morelos, sand dunes, VII-5-77 (1); 10 mi S Punta Prieta, VI-21-1938 (1); Rancho Mesquital, VI- 14/ 15-1967 (1); 6.2 mi NE Rancho Rosarito, VII-10-1979 (3); 6 mi S Rancho Rosarito, VIII-4-1977 (1); 1.6 mi W Rancho Rosarito, VI- 13/ 14- 1967(1). Eusattus dubius setosus, new subspecies Epistomum moderately to deeply and narrow- ly emarginate (as in Fig. 59), weakly indented at lateral sutures; antennal segment length ratios as follows: 2.5:1.0:2.2:1.6:1.6:1.4:1.4:1.4:1.4:1.2: 1 .6; terminal segment about 2 x as long as broad, apex attenuate or narrowly rounded; elytra set with muricate punctures about as large as eye DOYEN: SYSTEMATICS OF ELSATTL'S AND CONISATTUS 83 Figures 122-123. Posterior aspect of forelegs. 122. Eusattus pallidas pallidas. 123. E. p. adustus. facets anteriorly, becoming finely tuberculate lat- erally and on declivity; set with fine short to moderately long setae, especially evident later- ally and on declivity. Measurements. — EL 5.9-7.9 mm; EW 4.2- 5.6 mm; PL 1.9-2.3 mm; PW 3.8-5.1 mm; BD 3.2-4.4 mm. Holotype Female (CAS) and 3 paratypes (EME) from Mexico, Baja California Sur, Mu- lege, II-5- 1 974, G. Mckibbon; 2 paratypes (RLA) from 8.3 km SE Mulege, 111-26-1975, R. Aalbu, beach dunes. Diagnosis. — Eusattus d. setosus is distin- guished by the combination of deeply emarginate epistomum and relatively long 3rd antennal seg- ment (see Measurements). Distribution (Fig. 120). — Known only from the coastal sand dunes south of Mulege. Eusattus pallidus, new species (Figure 121) Very strongly convex, subglobular beetles with pale tan to chestnut, glabrate cuticle. Male. — Head and pronotum with simple punctate or tuberculate sculpturing; epistomum entire or weakly indented at lateral sutures, lat- eral lobes as wide as eyes; antenna subfiliform, with segments 9-1 1 enlarged as weak club; ter- minal segment 1. 1—1.2 x longer than broad; mentum 1.5-1.7 x broader than long, flat, with lateral borders arcuate, slightly raised in poste- rior third; anterior border shallowly, evenly emarginate; gula narrowed to V4-V3 width of sub- mentum. Pronotum 2.8-4.0 x as broad as long, lateral margins narrowly explanate, briefly upturned; disk with duplex punctation; presternum and prosternal process granulose. Elytra 1.0-1.25 x longer than broad, punctate or tuberculate; epipleuron set with long, mod- erately dense setae in basal third, glabrate api- cally; mesosternum between coxae slightly broader than prosternal process, not excavate; metasternum sparsely, setigerously punctate; se- tae short medially, long laterally; metasternal su- ture about Vi length metasternum. Protibia with row of spinules extending '/2— 2A distance to apex along outer margin (Figs. 122, 123); spinules separated by about 1 spine di- ameter; anterior face with scattered, short setae; metafemur with few long, projecting setae dor- sally or subglabrous; meso- and metatibia spi- nose; tarsomere length ratios as follows: 2.5:0.7: 0.6:0.5:2.0 (pro-); 3.5:1 .2: 1 .0:0.8:2.6 (meso-); 4.8: 1.6:1.3:3.2 (meta-). Female. — Spermathecal tubes 4, long slender; coxite length ratio to paraproct length = 0.26. Holotype Female (CAS) from Mexico, Baja California Sur, Isla San Jose, Bahia Amortajada, IV-1-1974, J. Doyen (J. Doyen Lot 74C1 1). Diagnosis.— Eusattus pallidus is distinctive in its small size, inflated, globular shape, and pale, often semitranslucent cuticle. It is most similar to E. vizcainensis Doyen, differing as stated in the diagnosis for the latter. It differs from E. ciliatus Horn, E. ciliatoides Doyen, and E. are- narius Doyen in its abbreviated row of protibial spinules. Variation.— There is significant geographic variation in size, color, cuticular sculpturing, and protibial armature. This variation is recognized here at the subspecific level. 84 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figure 124. Known distribution of Eusattus pallidus and E. vizcainensis. Key to the Subspecies of Eusattus pallidus 1. Cranium tuberculate; elytra muricately punctate; EW/EL < 0.86 2 Cranium and elytra very finely, obscure- ly punctate; EW/EL > 0.95 pallidus immaculatus 2( 1 ). Protibia bearing comb of 5-7 stout spines along posterior margin (Fig. 122) pallidus pallidus Protibia bearing row of long, slender se- tae alternating with short, stout spines (Fig. 123) pallidus adustus Eusattus pallidus pallidus, new subspecies Frons set with rounded tubercles V2-I eye facet in diameter, separated by 1 -3 tubercle diameters, becoming denser laterally and on epistomum; antennal segment length ratios approximately as follows: 1.0:0.6:1.0:0.7:0.8:0.7:0.7:0.6:0.6:0.6: 0.9. Pronotum 2.6-2.8 x broader than long; disk set with larger punctures V4-V2 eye facet in di- ameter, often obscured medially, and near lateral margins bearing flattened, yellow setae about as long as 3rd antennal segment; smaller punctures barely visible at 50 x. Elytra 1. 1-1.4 x longer than broad, pale tan or occasionally dark brown; disk set with muricate punctures about as large as eye facets, bearing short, declined setae, most evident laterally. Protibia bearing row of 5-7 short, stout spines along posterior margin (Fig. 122); apical protibial process extending to apex of 1st or 2nd tarsomere. Measurements. — EL 4.1-5.8 mm; EW 3.2- 4.3 mm; PL 1.0-1.4 mm; PW 2.6-3.8 mm; BD 2.7-3.7 mm. Holotype Female (CAS) and 41 paratypes (EME) from Mexico, Baja California Sur, Isla San Jose, Bahia Amortajada, IV- 1 - 1 974, J. Doy- en (J. Doyen Lot 74C1 1). Additional paratypes from Mexico, Baja California Sur, Isla Espiritu Santo, Bahia San Gabriel, III-3 1 - 1 974, J. Doyen (J. Doyen Lot 74C8) (28, EME); La Paz, X-18- 1974, W. W. Middlekauff (6, EME); 1-23-1974, 40 m, E. L. Sleeper (15, EME, CSULB). Distribution (Fig. 124) and Habitat.— Eu- sattus p. pallidus occurs in coastal sand dunes on the shores of the Gulf of California from La Paz north to Bahia Evaristo, and on several gulf is- lands. Adults are abundant beneath strand and dune vegetation, especially Atriplexsp., from just above the reach of highest tides to 25-40 ft be- hind the beach, where they are associated with Cryptadius sinuatus Blaisdell. Additional Material Examined. — Mexico: Baja Califor- nia Sur: 2 mi E El Coyote, NE La Paz. XII-30-1958, near ocean beach [corpses] (10); Bahia Evaristo, IV-5-1974 (3); south end Isla San Francisco, IV- 1 1/12-1974 (3); landlocked lagoon. Nend Isla San Jose, IV-3-1974 (1). Eusattus pallidus adustus, new subspecies Similar to E. p. pallidus, except in the follow- ing features: Frons with tubercles separated by 1-4 diameters, becoming obscured posteriorly between eyes. Elytra dark brown, disk set with strongly muricate punctures or weak tubercles about '/2 eye facet diameter. Protibia bearing row of short, stout spines alternating with long slen- der setae along posterior margin (Fig. 123). Measurements. — EL 5.0-6.4 mm; EW 4.0- 5.0 mm; PL 1.3-1.6 mm; PW 3.5-4.5 mm; BD 3.4-4.0 mm. Holotype Male (CAS) and 22 paratypes DOYEN: SYSTEMATICS OF EUSATTUS AND CONISATTUS 85 (EME, RLA) from Mexico, Baja California Sur, San Bruno, 14 mi SE Santa Rosalia, VII-7-1979, R. L. Aalbu; 18 paratypes (EME, CDFA), same data. Hardy, Andrews, Giuliani; 2 paratypes (CDFA), 13 mi SW Guillermo Prieto, 111-31- 1982, M. Wasbauer. Diagnosis.— Eusattus p. adust us is of larger average body size than E. p. pallidas, is usually darker in color, and differs in its protibial spi- nation (Figs. 122-123). It is known only from beach dunes at the type locality (Fig. 1 24), where the beetles were sifted from sand beneath shrubs about 100 m inland (A. Hardy, pers. comm.). Eusattus pallidas immaculatus, new subspecies Similar to E. pallidus pallidas, differing as fol- lows: Frons glabrous or with few very fine punc- tures near epistomal suture, denser and slightly larger on epistomum. Pronotum 2.8-2.9 x broader than long; disk set with larger punctures about Va eye facet diameter, bearing very short, fine setae near lateral margins; smaller punctures about xli size of larger. Elytra about 1.05 x longer than broad, set with punctures about 'A eye facet diameter, surrounded by slight depressions and separated by 6-10 puncture diameters; elytral setae exceedingly fine and short, visible only un- der oblique lighting. Protibia with row of short, stout spines alternating with long, slender spines along posterior margin. Measurements. — EL 3.6-4.3 mm; EW 3.4- 4.1 mm; PL 1.0-1.3 mm; PW 2.9-3.6 mm; BD 3.0-3.4 mm. Holotype Female (CAS) and 3 paratypes (EME, CSULB) from Mexico, Baja California Sur, 47 km SE Guerrero Negro, 1-15/16-1974, 240 m, E. L. Sleeper. Diagnosis.— Eusattus p. immaculatus is dis- tinguished by its very finely sculptured cuticle. It is known only from the type locality (Fig. 1 24). Eusattus vizcainensis, new species Strongly convex, ovoid beetles with dark brown to black, glabrate cuticle. Male.— Frons set with rounded tubercles about as large as eye facets, separated by 1-3 tubercle diameters, becoming denser anteriorly and sub- contiguous laterally near eyes; epistomum entire or weakly indented at lateral sutures; lateral lobes about as prominent as eyes. Antennal segment length ratios as follows: 1.4:0.7:1.4:1.0:0.9:0.9: 0.8:0.7:0.8:0.7:0.9; terminal segment about 1.1 x Figure 125. Posterior aspect of left foreleg of Eusattus vizcainensis. longer than broad, apex rounded or obliquely truncate. Mentum about 1 .6 x broader than long, anterior margin shallowly and evenly concave; gula narrowed to V5-V6 width of submentum. Pronotum 2.7-3.0 x wider than long, widest about 3A distance from apex to base; disk set with punctures about lh eye facet diameter, separated by 2-4 puncture diameters, becoming coarser, closer near lateral margin, and bearing setae about lh length 2nd antennal segment; lateral margins narrowly explanate and subhorizontal; prester- num granulose, prosternal process more finely so. Elytra 1.2-1.3 x longer than broad, set with rounded tubercles about as large as eye facets, separated by 2-4 puncture diameters; tubercles supertending very short, declined setae; epipleu- ron sparsely setose in basal third, glabrate api- cally. Mesosternum between coxae broader than prosternal process, not excavate. Metasternum glabrate medially, set with moderately coarse, setigerous punctures laterally. Protibia with api- cal process narrowly rounded, extending as far as apex of 1 st tarsomere; outer margin with row of spinules separated by 1-4 spinule widths ex- tending to apex (Fig. 125); posterior margin bear- ing fringe of long, projecting setae, sometimes interspersed with 1 or 2 shorter, spinose setae. Metafemur with short, declined setae; meso- and metatibiae spinose; tarsomere length ratios as follows: 1 .4:0.5:0.5:0.4:2.0(pro-); 2.9: 1.2:1.1 :0.8: 2.5 (meso-); 4.2:1.5:1.2:3.3 (meta-). Female. — Spermathecal tubes 2, long, slender; coxite length ratio to paraproct length = 0.25. Measurements. -EL 3.8-5.2 mm; EW 3.1- 3.9 mm; PL 1.0-1.1 mm; PW 3.0-3.7 mm; BD 2.7-3.3 mm. Holotype Female (CAS) and 2 paratypes (CDFA) from Mexico, Baja California Norte, 6 86 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 126 Figures 126-127. Eusattus dcpressus. 126. Ovipositor, ventral (left) and lateral (right) aspects. 127. Posterior aspect of left foretibia. mi N Guerrero Negro, VIM- 1979, Hardy, An- drews, and Giuliani. 5 paratypes (EME) from Baja California Sur, 7 mi SE Guerrero Negro, IV-8-1976, J. Doyen, P. Rude, R. Morrison. Diagnosis.— Eusattus vizcainensis is similar to E. palhdus Doyen, differing in having the row of spinules extending the entire length of the out- er tibial margin (extending xli-2k length in palh- dus). The elytral tubercles of vizcainensis are much coarser than those of pallidus, and the cuticle is very dark brown or black (pale tan to chestnut in pallidus). E. vizcainensis is less globose than pallidus. E. vizcainensis resembles small indi- viduals of E. ciliatoides Doyen, but the anterior tibial spinules of the latter are contiguous or nearly so (separated by 1-4 spine widths in vizcainen- sis), the frons is punctate (tuberculate in vizcai- nensis), and the anterior margin of the mentum is nearly straight (evenly concave in vizcainen- sis). Distribution (Fig. 124) and Habitat.— Sand dunes in the western Vizcaino Desert. Eusattus depressus Species Group Moderately convex, black beetles with pro- sternum expanded anteriorly to cover gular re- gion; intercoxal process obtusely angulate. Head and pronotum with simple or duplex punctation; epistomum with very shallow, nar- row medial emargination (Fig. 61), weakly in- dented or concave at lateral sutures; epistomal suture obscured by cranial sculpturing; pyriform or very weakly reniform, barely indented by epi- stomal canthus, dorsal lobe much broader than ventral; antennae extending to base of pronotum or slightly beyond, scape subglabrous, flagellum pubescent, gradually enlarged to 10th segment, submoniliform; mentum nearly flat, coarsely punctate, lateral margins arcuate, anterior angles rounded; submentum subcrescentic, slightly nar- rower than base of mentum; gular sutures strong- ly convergent, separated by Kk-Xk submentum width anteriorly. Pronotum strongly convex in lateral silhou- ette, discontinuous with contour of elytra; lateral margins very briefly or not explanate; hypome- ron glabrous or setose; prosternum prolonged an- terad of coxae, longer than prosternal process and concealing mentum in deflexed head (Fig. 45); prosternal process at least % as wide between coxae as behind, broadly rounded. Elytra with epipleuron gradually narrowing from base to apex; pseudepipleuron rounded, un- defined; mesosternum subequal in width to me- sosternal process, deeply excavate; median meta- sternal suture absent; intercoxal process broadly rounded or obtusely angulate with broadly rounded apex (Fig. 43). Legs variable. Metendosternite with arms fused with meso- coxal inflections, fused apically with meso- notum; mescndosternite with arms strongly ex- panded basally as muscle attachment disks; spermathecal tubes 2, multiply branched, long, slender, convoluted (Fig. 9); aedeagus variable; ovipositor subglabrous, paraproct expanded ventrally beneath coxite (Fig. 1 26); spatulate pro- cesses of coxites subparallel, bluntly rounded. DOYEN: SYSTEMATICS OF EUSATTUS AND COS IS A I 77 .S 87 upcurved; coxite length ratio to paraproct length a 0.34. Eusattus depressus Champion Eusattus depressus Champion, 1884:75, table 4. Eusattus puncticeps Blaisdell, 1923:269, new synonymy. Eusattodes depressus, Gebien 1938:283 (402). Oval, dull black, somewhat flattened beetles. Male. — Frons punctato-rugose or punctate, larger punctures V/i-2 eye facets in diameter, subcontiguous, or separated by 1-3 puncture di- ameters, becoming coarser, closer near epistomal suture; smaller punctures xk eye facet diameter or less, separated by about 1 puncture diameter; epistomum punctato-rugulose, margins not re- flexed; medial epistomal suture weakly im- pressed, sometimes partly obscured by cranial sculpturing; lateral sutures faint, usually ob- scured. Antennal segment ratios as follows: 3.7: 2.3:3.2:2.6:2.6:2.6:2.4:2.4:2.0: 1 .9:2.6; terminal segment about 1 .3 x longer than broad, apex ob- tusely angulate; mentum about 1 .5 x broader than long, anterior border evenly and moderately emarginate. Pronotum 1.8-2.0 x broader than long, lateral margins nearly straight in posterior half, then evenly convergent to nearly right-angled anterior corners; posterior corners acute, strongly pro- duced posteriorly, overlapping humeral angles; posterior border weakly arcuate or nearly straight between corners; disk set with punctures about '/4-V2 eye facet diameter, separated by 2-4 punc- ture diameters, becoming coarser and closer lat- erally; hypomeron longitudinally, somewhat sin- uously furrowed, glabrous except for few very short setae just below lateral margin; presternum punctato-rugulose, sparsely set with short, de- clined setae; prosternal process punctato-rugu- lose, becoming coarsely punctate or almost smooth posteriorly, distinctly margined or oc- casionally unmargined. Elytra 1.4-1.5 x longer than broad, lateral margins subparallel in anterior half, then con- verging to apex; disk alutaceous, set with punc- tures about xh eye facet diameter, separated by 1-4 puncture diameters; epipleuron glabrous; in- tercoxal process obtusely angulate. Femora with short, sparse, declined setae; pro- tibia narrowly explanate (Fig. 127), outer margin concave or nearly straight with row of spinules extending %-7/8 distance to apex, separated by 2- 4 spinule widths (Fig. 127); apical process ex- tending about lh distance to apex of basal tar- somere; posterior face coarsely punctate basally, finely, closely punctate or punctato-granulose apically; meso- and mctatibiae spinose; protar- sus subequal in length to protibia; tarsomere length ratios as follows: 5.5:2.5:2.2:2.0:6.7 (pro-); 6.4:3.0:2.6:2.0:6.0 (meso-); 8.8:4.0:3.5:7.0 (meta-). Aedeagus and median lobe sharply attenuate (Fig. 53). Mesendosternite with arms extending about xh distance to elytral articulations. Female.— Apical protibial process extending to apex of 1 st or sometimes 2nd tarsomere; pro- tarsus % length of foretibia, tarsomere length ra- tios: 4.0: 1.5: 1 .4: 1 .3:4.2; otherwise differs as stat- ed in generic description. Measurements. — EL 8.5-1 1.0 mm; EW 5.8- 8.4 mm; PL 3.0-4.3 mm; PW 5.5-8.4 mm; BD 4.6-6.1 mm. Holotype Female in BMNH. Type Localities.— Of depressus, Mexico; Ala- mos, Sonora; puncticeps, San Pedro Bay, Sonora. Diagnosis.— Eusattus depressus is closely re- lated only to E. secutus Horn, but is much larger; has the anterior tibiae narrowly explanate, with spinose posterior face (broadly so with granulose posterior face in secutus); and has the hypomeron subglabrous (entirely setose in secutus). It is su- perficially similar to laevis LeConte, but the epi- pleuron is gradually narrowed (abruptly nar- rowed just behind humerus in laevis). Distribution (Fig. 1 28) and Habitat. — Thorn forest, especially riparian situations, from sea level to 4000' elevation, from southern Nayarit to northeastern Sonora. Adult activity is strongly concentrated in the summer rainy period from late June to mid-September. Additional Material Examined. — Mexico: Chihuahua: La Bufa, Sierra Madre Mtns., 900 m, VII-7-1 972 (1). Nayarit: 24 mi SE Tepic, VIII- 16- 1960 (2). Sinaloa: 4 mi NW Choix, VII- 17-1968 (1); 20 mi SE El Fuerte, VII-12-1962 (1); nr. Imla, E ofCuliacan, 457 m, X-22-1973 (1); Los Mochis, VIII-17-1922 (1); 4 mi N Piaxtla, VIII-1-1965 (1); Otates, V-25-1956 (1). Sonora: 3.7 mi SW La Aduana, IX-5- 1 964 ( 1 ); 1 5 mi W Agia- bampo, 1 00', I V-29- 1 949 ( 1 ); Alamos and vicinity, VI- 1 8/IX- 19 (80); Guirocoba, VH-5-1933 (1); 3 mi N Hermosillo, V-25- 1961 (1); 4 mi W Mazatlan, VIII-17-1964 (1); 10 mi NW Mazocahui, 3800', 111-26-1980 (2 corpses); 1 1 mi S C. Obre- gon, Vlll-l 1-1960 (14); San Bernardo (Rio Mayo), VIII-21- 1935 (9); San Carlos Bay, VIII-10-1960 (1), IV-19-1974 (1). Eusattus secutus Horn (Figure 129) Eusattus secutus Horn, 1894:349, 421, 423; Casey 1908:65; Blaisdell 1943:193. Convex, oval or slightly pyriform, shining black beetles. Male. — Frons punctato-rugose, punctures 1- 88 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 DEPRESSUS O SECUTUS prosternal process, sparsely set with short, ap- pressed setae; prosternal process margined. Elytra 1.2-1.3 x longer than broad, lateral margins arcuate from base to apex; disk shining, faintly and sparsely rugulose, set with punctures about xk-lh eye facet diameter, separated by 2- 6 puncture diameters; epipleuron sparsely setose in basal third; intercoxal process broadly round- ed. Profemur with dorsal surface sparsely set with long, projecting setae; protibia broadly expla- nate, subtriangular (Fig. 131), outer margin con- vex, reflexed posteriorly, with row of spinules extending about Vi distance to apex, separated by 1-4 spinule widths; apical process broadly to narrowly rounded, extending to apex of 1st or 2nd tarsomere; posterior face finely punctato- granulose; mesofemur sparsely set with moder- ately long, inclined setae on dorsal surface; meta- femur subglabrous; tibiae spinose; protarsus about % length of tibia; tarsomere length ratios as follows: 2.9: 1 . 1 :0.9:0.8:2.6 (pro-); 3.4: 1.5:1.3: 1.0:2.7 (meso-); 5.3:1.9:1.6:3.3 (meta-). Aedea- gus with apex truncate, median lobe attenuate (Fig. 1 30). Mesendosternite with arms extending about % distance to elytral articulations. Figure 128. E. secutus. Known distribution of Eusattus depressus and 1.5 eye facets in diameter, separated by 1-2 puncture diameters, becoming more strongly ru- gose on epistomum; epistomal margins briefly reflexed; epistomal suture weakly impressed or obscured by cranial sculpturing. Antennal seg- ment ratios as follows: 2.2: 1.5:1.9:1 .6: 1 .6: 1 .6: 1 .6: 1.6:1.5:1.4:2.0; terminal segment about 1.75 x longer than broad, ovoid, with subangulate apex near 90°; mentum about 1 .5 x broader than long, anterior border angulately emarginate. Pronotum 1 .7—1.9 x broader than long, lateral margins evenly arcuate in anterior 3/4-%, then nearly straight to posterior angles; anterior angles nearly 90°, posterior angles narrowly acute, strongly produced posteriorly, contiguous with elytral humeri; disk shining, set with punctures about xk-xh eye facet diameter, separated by 2- 4 puncture diameters; hypomeron unfurrowed except over coxae, sparsely set with setae, longest medially; prosternum with anterior margin strongly elevated; smooth laterally, becoming coarsely punctato-rugulose medially and on Figure 129. Eusattus secutus Horn. DOYEN: SYSTEM ATICS OF EUSATTUS AND COMSAT 1 1 s 89 130 V Figures 1 30-1 3 1 . Eusattw secutus Horn. 1 30. Tegmen (left) and median lobe (right). 131. Posterior aspect of left foreleg. Female. -Differs as stated in generic and species group descriptions. Measurements. -EL 6.5-8.3 mm; EW 4.8- 6.4 mm; PL 2.5-3.3 mm; PW 4.4-5.7 mm; BD 4.0-5.1 mm. Lectotype Female (?) (CAS) from Mexico, Baja California Sur, El Taste; 1 syntype (missing) from Cabo San Lucas. Diagnosis.— Eusattus secutus is closely relat- ed only to E. depressus Champion, differing as stated in the diagnosis for the latter. Superficially secutus resembles E. dubius LeConte, but has the epistomum very shallowly emarginate (moder- ately to deeply emarginate in dubius), and also differs from dubius in the characters listed in the key to subgenera. Except for Conisattus rectus Casey, secutus is unique among Eusatti in having the entire hypomeron setose. Distribution (Fig. 128) and Habitat.— This seldom-collected species occurs in thorn-forest habitats from San Jose Comondu south to Cabo San Lucas, at elevations from near sea level to at least 300 m. Additional Material Examined. — Mexico: Baja Califor- nia Sur: 3.3 mi S El Cien, IX-26-1981 (3); 21 mi W La Paz, VIII-9-1966 (3); 26 mi W La Paz, VIII-10-1966 (1); 59 mi NW La Paz, X-24-1968, 600 ft, (2); 8 mi W La Paz, 1000 ft, X-3-1968 (1), X-13-1968 (1); Las Cuevas, 111-25-1975 (1 corpse): Rancho El Palomar, X-20-1972 (3); 15 km SW San Jose Comondu, 111-22-1975; San Hilario, 1000', XI-5-1968 (19); 3 mi N Santiago, VII-16-1957 (1); Sierra El Chinche (2); Todos Santos, X- 1 0- 1 94 1 (2); 6 mi E Todos Santos, X-4- 1 98 1 (2); 9.3 mi SE San Perdito, X-7-1981 (1). Eusattus rudei Species Group Oval, moderately convex beetles with finely sculpted, shining cuticle. Head and pronotum with duplex punctation; eyes ovoid or very weakly reniform, scarcely in- dented by epistomal canthus, dorsal and ventral lobes subequal; antenna extending about 2h dis- tance to base of pronotum, submoniliform, grad- ually enlarged to 10th segment, scape and fla- gellum pubescent; mentum punctate, set with long, projecting setae, lateral borders arcuate, an- terior border moderately deeply emarginate, an- terior angles acutely rounded; submentum sub- equal in width to base of mentum; gular sutures gradually converging, separated by about Vi-Va mentum width anteriorly. Pronotum 2. 1-2.3 x broader than long, disk with duplex punctation, strongly convex in trans- verse section, evenly declivous to lateral mar- gins, narrowly beaded except posteriorly, streamline with elytra in longitudinal silhou- ette; broadest at base, lateral margins evenly ar- cuate, anterior angles obtuse, posterior angles acute; hypomeron glabrous, polished except for dense fringe of submarginal setae projecting well beyond pronotal borders and few shorter setae anteromedially; presternum before coxae much shorter than prosternal process, sparsely to densely and setigerously punctate, setae long, slender, and erect; prosternal process broadly rounded, unmargined, slightly more than lh as wide between as behind coxae, densely and finely 90 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figures 132-133. Posterior aspect of left forelegs. 132. E. crypticus. 133. E. rudei. punctate, with few large punctures and setae su- perimposed. Elytra 1.3-1 .4 x longer than broad, with lateral margins arcuate; pseudepipleura broadly round- ed, undefined; epipleuron gradually narrowing from base to apex, finely punctate and setose along basal Vi-Va of outer margin; mesosternum between coxae subequal in width to prosternal process, barely excavate; metasternum moder- ately coarsely punctate, sparsely setose; median metasternal suture about lh length metasternum; intercoxal process subtriangular, acutely round- ed (as in Fig. 42). Anterior and middle femora bearing dense fringes of long erect or retrorse setae on dorsal surfaces; foretibia with spinule row extending Vi distance or slightly less to apex, spinules sepa- rated by 1-3 spine widths (Figs. 132, 133); mid- dle and posterior tibiae spinose. Metendosternite with arms fused with meso- notum, not fused with mesocoxal inflections; mesendosternite with arms extending about Vi distance to elytral articulations, expanded ba- sally; spermathecal tubes 4, unbranched or pau- cibranched, long, slender, tightly convoluted; va- gina with large, pleated proximal bursa (Fig. 10). Aedeagus and parameres with apices bluntly rounded; spatulate processes of coxites bluntly rounded (Fig. 15), not upcurved; coxite length ratio to paraproct length variable. Eusattus crypticus, new species Very dark brown to black beetles with tuber- culate elytral declivity. Male. — Head amplected to anterior margin of eyes when retracted; frons set with punctures about 1-1 V2 eye facets in diameter, separated by about 1-3 puncture diameters; smaller punctures about '/3-'/4 eye facet diameter, subcontiguous or separated by about 1 diameter; epistomum deep- ly and narrowly emarginate medially (as in Fig. 59), barely indented or entire at lateral sutures; epistomal suture usually obscured by sculptur- ing, especially medially. Antennal segment ratios as follows: 2.2:1.0:1.5:0.9:1.0:1.0:1.0:1.0:1.0:0.8: 1.1; terminal segment slightly broader than long, apically rounded. Mentum flat, deeply grooved along posterior half of lateral margins; submen- tum slightly narrower than base of mentum. Pronotal disk set with larger punctures about V2 eye facet diameter, separated by about 1-3 puncture diameters; smaller punctures barely visible at 50 x . Elytral disk finely, sparsely crin- kled; set with punctures about Vi eye facet di- ameter, separated by about 3-6 puncture di- ameters anteriorly, becoming coarser and closer and distinctly muricate in middle of disk, grad- ually transforming into tubercles about 2 eye fac- ets in diameter on declivity. Anterior tibia with sinuous outer margin, more abruptly enlarged near apex (Fig. 132); apical process extending approximately to apex of 2nd tarsomere; anterior face set near base with sev- eral long, slender setae; posterior face with 4-5 spines in basal fourth or fifth, glabrous, polished apically; apical border without spines internally, bearing 4-5 spinules externally near tibial spurs; posterior femur bearing sparse fringe of long se- tae on dorsal surface; tarsomere length ratios as follows: 3.5:0.8:0.8:0.7:2.2 (pro-); 4. 1 : 1 .7: 1 .5: 1.3: 2.7 (meso-); 5.8:2.1:1.7:3.5 (meta-). Female.— Coxite to paraproct ratio = 0.52 (Fig. 15); otherwise differs as stated in generic description. Measurements. — EL 6.6-8.4 mm; EW 5.1- DOYEN: SYSTEMATICS OF Ei'SATTUS AND COMSATTi'S 91 6.5 mm; PL 2.3-2.8 mm; PW 4.9-6.0 mm; BD 4.0-5.0 mm. Holotype Female (CAS) and 8 paratypes (EME, CSULB) from Mexico, Baja California Sur, Los Frailes, IV-25/26-1975, E. M. Fisher; 4 paratypes (CAS) from Bahia de Los Frailes, III- 19- 1953, J. P. Figg-Hoblyn; 17 paratypes (EME) from Las Barracas, 30 km E La Ribera, III-2 1/24- 1982, coastal dunes, E. I. Schlingerand M. E. Irwin; 8 paratypes (CDFA), Los Barriles, X-3-1981, under logs on beach, D. Faulkner, F. Andrews. Diagnosis. —Eusattus crypticus is similar only to E. rudei Doyen, differing as described in the diagnosis for the latter. Distribution (Fig. 134) and Habitat.— This species is presently known only from the ex- tremely arid coast east of the Sierra el Trinidado. The beetles burrow in the sand about the bases of plants on foredunes a few meters above sea level (M. E. Irwin, pers. comra.). Eusattus rudei Doyen Black beetles with alutaceous or finely rugulose elytra. Male. — Head amplected to posterior margin of eyes; frons set with larger punctures 1-1 lh eye facets in diameter, separated by 1-2 puncture diameters; smaller punctures barely visible at 50 x, subcontiguous; epistomum shallowly emarginate medially (as in Fig. 61), moderately indented at lateral sutures; epistomal suture usu- ally partly obscured by cranial sculpturing: an- tennal segment ratios as follows: 3.3:1.4:2.2:1.5: 1.6:1.7:1.5:1.5:1.5:1.5:1.4:1.6; terminal segment about as wide as long, subtriangular with round- ed apex. Mentum slightly convex, lateral borders reflexed in posterior half; submentum slightly wider than base of mentum. Pronotal disk set with larger punctures lU-lh eye facet diameter, separated by 2-6 puncture diameters; smaller punctures barely visible at 50 x . Elytral disk finely, sparsely crinkled ante- riorly, becoming rugulose on declivity; puncta- tion duplex; larger punctures '/2-1 eye facet in diameter, separated by 2-6 puncture diameters anteriorly, becoming coarser and closer on de- clivity; smaller punctures obscured, most appar- ent on declivity. Anterior tibia subtriangular (Fig. 133), apical process extending about to apex of 1 st tarsomere; anterior face with few spines and short, decum- • RUDEI O CRYPTICUS Figure 1 34. Known distribution of Eusattus crypticus and E. rudei in the Cape region of Baja California. bent setae; posterior face with basal lh spinose, apical % glabrous, polished, apical border mar- gined with spinules on both anterior and pos- terior edges; posterior femur bearing few. mod- erately long, decumbent setae; tarsal length ratios as follows: 5.0:1.1:1.1:1.0:3.6 (pro-); 6.8:2.3:2.2: 1.9:4.9 (meso-); 8.2:3.0:2.3:5.0 (meta-). Female. — Coxite to paraproct ratio = 0.37; otherwise differs as stated in generic and species group descriptions. Measurements. — EL 8.7-1 1.4 mm; EW 6.4- 8.5 mm; PL 2.7-3.7 mm; PW 6.0-8.3 mm; BD 5.0-6.8 mm. Holotype Female (CAS) and 26 paratypes (EME) from Mexico, Baja California Sur, Cabo Falso, 7 km W Cabo San Lucas, 1-1-1979, P. Rude, on sand dunes; additional paratypes: Cabo San Lucas, Pacific side, XII-8-1979, L. G. Frei- hofer ( 1 , CAS); 5 . 5 mi NW Todos Santos on road to la Pastora, 1-13-1959, H. B. Leech (1, CAS); 4 mi S El Pescadero, X-24-1968, E. L. Sleeper (1, CSULB); 18.8 mi S Todos Santos, IX-29- 1981, beach dunes, D. Faulkner, F. Andrews (30, EME, CDFA). Diagnosis.— Eusattus rudei is closely related only to E. crypticus Doyen, but is much larger, has the elytral declivity rugulose rather than tu- berculate, and has the outer foretibial margin nearly straight (see Figs. 132, 133). It is super- ficially similar to E. laevis LeConte, but has the thoracic venter densely setose (glabrous in laevis) and the epipleuron gradually narrowed (abruptly narrowed behind humerus in laevis). Distribution (Fig. 134) and Habitat.— Known onlv from maritime sand dunes along 92 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Figures 135-136. Posterior aspect of left forelegs. 135. Eusattus politus. 136. E. robustus. the west coast of extreme southern Baja Califor- nia. Eusattus robustus Species Group Head and pronotum with duplex punctation; epistomum moderately to deeply and angulately emarginate medially (as in Fig. 59), scarcely or not indented at lateral sutures; eyes weakly re- niform, dorsal and ventral lobes subequal; an- tenna setose, subfiliform basally, then gradually broadened to apex; terminal segment 1.1-1.3 x broader than long, symmetrical, with truncately rounded apex; mentum convex, lateral margins arcuate, more convergent basally; submentum as wide as base of mentum. Pronotum strongly convex, broadly explanate laterally, broadest just before base; lateral mar- gins arcuate; anterior corners nearly 90°, round- ed; antecoxal part of presternum much shorter than prosternal process; prosternal process about 2x as wide behind as between coxae, slightly convex, margined, broadly rounded, semicir- cular posteriorly. Elytra strongly convex, smooth or weakly alu- taceous; pseudepipleuron rounded, undefined, or absent; epipleuron gradually narrowed from base to apex; mesosternum shallowly excavate; meta- sternum finely punctate, laterally setose; meta- sternal suture about lh length metasternum. Meso- and metafemora with sparse, dorsal fringes of long, projecting setae; protibia broadly triangular, with attenuate apical process extend- ing to apex of 2nd or 3rd tarsomere (Figs. 135, 1 36); outer margin with row of closely spaced or subcontiguous spinules extending to apex; meso- and metatibiae clavate, more abruptly expanded near apex. Metendosternite with apices of arms fused with metanotum, not fused with mesocoxal inflec- tions; mesendosternite with arms extending about '/2 distance to mesonotum, flanged basally. Ae- deagus bluntly rounded; spermathecal tubes 4, long, slender, unbranched (Fig. 11). Spatulate process of ovipositor straight; coxite length ratio to paraproct length variable. Eusattus politus Horn Eusattus politus Horn, 1883:304. Convex, broadly oval, shining black beetles. Male. — Head and pronotal disk with duplex punctation, epistomum deeply emarginate (as in Fig. 59); antennal segment ratios as follows: 3. 1: 1 .0: 1 .5: 1 . 1 : 1 .0:0.9:0.9:0.9: 1 .0: 1 .0: 1 .0; mentum wth anterior corners nearly right-angled, round- ed; anterior margin sinuous, deeply emarginate; gular sutures confluent in anterior third. Pronotum 2.2-2.6 x broader than long, widest V4-V6 before base, sides evenly arcuate; posterior corners slightly acute, briefly produced poste- riorly; disk with large punctures separated by 2- 4 diameters medially, 1-2 diameters laterally; small punctures evenly distributed 1-2 diame- ters apart; hypomeron smooth, shining, bearing marginal fringe of long, projecting setae and scat- tered long setae centrally; sternum scabrous, opaque, sparsely set with long projecting or ap- pressed setae; prosternal process with few de- clined setae anteriorly. Elytra 1.2-1.3 x longer than broad; setigerous near epipleuron; pseudepipleural margin broadly rounded; epipleuron narrowing to half basal width by 1st abdominal sternite, then gradually nar- rowing to apex. Mesosternum weakly excavate between coxae, % width of prosternal process. DOYEN: SYSTEMATICS OF ELSATTLS AND ( ONIS I III S 93 Profemur set with sparse, moderately long pro- jecting setae dorsally (Fig. 135); mesotibia and metatibia with anterior and ventral surface clothed with long erect or retrorse setae, with shaft set with blunt spines ventrally, spinose se- tae dorsally; tarsomere length ratios: 2.9:1.0:0.8: 0.7:2.7 (pro-); 4.9: 1 .7: 1 .4: 1 . 1:3.0 (meso-); 7. 1 :2.3: 1.6:3.8 (meta-). Female. — Coxite length ratio to paraproct length = 0.28; otherwise differs as stated in ge- neric description. Lectotype Female (MCZ) from Santa Bar- bara, California. Diagnosis.— Eusattus politus is similar in body form to E. robustus LeConte, but much smaller in size. In politus, the narrower epipleuron does not reach the margin of the elytral disk; in ro- bustus, the epipleuron is coincident with the mar- gin. Variation.— The variation in cuticular sculp- turing and setation of this species is recognized here at the subspecific level. Eusattus politus politus Horn Eusattus politus Horn, 1883:304; 1894:423 (key); Fall 1897: 238 (distribution); 1901:166 (distribution); Casey 1908:68; Cockerell 1939:283. Eusattus vanduzeei Blaisdell, 1921:214; Cockerell 1939: 283, new synonymy. Eusattus (Eusattus) vanduzeei, Leng and Mutchler 1927:35. Eusattus vanduzei, Gebien 1938:284 (403); Papp 1961:116 (misspelling). Head and pronotum with large punctures sub- equal to eye facets in size, small punctures V4-V6 that size; frons with large punctures separated by about 1-3 puncture diameters, slightly denser near epistomal suture and on epistomum; small punctures separated by 1-2 puncture diameters. Elytral disk with larger punctures subequal to larger pronotal punctures, ranging to Vi that size, separated by 3-8 puncture diameters, setigerous near epipleuron; epipleuron sparsely set with long setae, becoming glabrous in posterior third. Pro- tibia with spine field occupying basal third of posterior face (Fig. 135). Measurements. — EL 5.7-8.8 mm; EW 6.2- 8.1 mm; PL 1.9-2.9 mm; PW 5.1-7.0 mm; BD 4.2-7.0 mm. Lectotype Female in MCZ. Type Localities. — Of politus, Santa Barbara, California; vanduzeei, Prince Island (near San Miguel Island), Santa Barbara Co., California. Diagnosis.— The more finely punctate, less se- tose dorsum distinguishes this subspecies from the next. Distribution and Habitat. — This subspecies is restricted to San Miguel and Santa Rosa is- lands and Prince Island, an islet near San Miguel. A few specimens have associated ecological in- formation suggesting that the preferred substrate is sand— but, as in the case of E. robustus, most habitats on the islands are probably occupied. Aside from the label data on a few specimens (including the type) collected before the turn of the century, there is no evidence that E. politus occurs anywhere on the mainland. Additional Material Examined. — California: Santa Bar- bara Co.: San Miguel Island: no further data (45); Cuyler Har- bor, VII- 11-1 980, VIII-3 1 - 1 978 (2); San Miguel Mtn.. VII- 1 1 - 1970 (1); Simington Cove, VII-1 1-1970, VIII-29-1978 (7); W end sand dunes, V-20- 1 977, V-23- 1 978 (7); Willow Creek, IV- 25-1979 (1); Prince Island: V- 19- 1919 (55); Santa Rosa Island: no further data, IX- 14- 1974 (4); Beechers Bay, XII-27/28- 1972 (1); Skunk Point Dunes, VII-2-1971 (5). Eusattus politus cruzensis, new subspecies Head and pronotal disk set with large punc- tures about 2 eye facets in diameter, separated by about xh-2 puncture diameters on frons, be- coming almost contiguous on epistomum; small punctures about Xh-Vi that size. Elytral disk with larger punctures muricate, setigerous, subequal to large pronotal punctures, separated by about 4-6 puncture diameters; setae suberect or de- clined, as long as hypomeron setae laterally, de- creasing to vh that length centrally; setigerous punctures interspersed with simple punctures about lh as large, separated by 2-4 diameters. Measurements. — EL 5.9-6.2 mm; EW 4.9- 5.4 mm; PL 2.0-2.1 mm; PW 4.7-5.2 mm; BD 3.5-4.0 mm. Holotype Female (CAS) from California, Santa Barbara County, Santa Cruz Island, Can- yon del Medio, V-9-1969, R. O. Schuster. Be- sides the type, a single specimen in the Horn Collection shows the characters of E. p. cruzen- sis; it bears the label "S. Barbara," as do Horn's specimens of E. p. politus. Because of its broken, abraded condition, this specimen is not desig- nated as paratype. Eusattus robustus LeConte (Figure 137) Eusattus robustus LeConte, 1866:1 12; Horn 1883:304; 1894: 423 (key); Fall 1897:238 (distribution); Doyen 1972:369 (morphology); 1974a:86 (predation); 1977:6 (synonymy). Nesostes robustus, Casey 1908:58; Cockerell 1939:283. 94 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 Figure 137. Eusaltus robustus LeContc. Nesostes robustus postremis Casey, 1908:59. Nesostes postremis, Cockerell 1939:283. Very broadly oval, moderately convex, black beetles with the epipleura occupying the entire inflexed part of the elytra. Male. — Head and pronotal disk with duplex punctation; frons with large punctures 1-2 eye facets in diameter, separated by 1-3 puncture diameters, becoming finer, sparser on vertex and near eyes, sparser and usually obscured near epi- stomal margin; smaller punctures about lA size of larger, separated by 2-4 puncture diameters; antennal segment length ratios as follows: 4.3: 1.7:2.6:2.5:2.5:2.4:2.3:2.3:1.8:1.7:1.7. Mentum with anterior margin shallowly, angulately emar- ginate, anterior angles acute, narrowly rounded; gula gradually convergent anteriorly, '/, width submentum. Pronotum 2.3-2.5 x broader than long; lateral margins evenly arcuate; posterior corners acute, strongly produced posteriorly; anterior angles nearly right-angled, narrowly rounded; disk with larger punctures about as large as eye facets, sep- arated by 1-4 puncture diameters; smaller punc- tures about VA that size, separated by 1-4 diam- eters; hypomeron shining, glabrous except for fringe of long setae along posterior half of lateral margin; sternum opaque, sparsely set with mur- icate punctures bearing moderately long, in- clined setae; prosternal process glabrous or near- ly so. Elytra 1.2-1.3 x longer than broad, lateral margins arcuate; disk alutaceous, set with tuber- cles 1-2 eye facets in diameter, separated by 1- 4 tubercle diameters, becoming slightly coarser laterally; on declivity more finely, sparsely tu- berculate and granulose; epipleuron occupying entire inflexed portion of elytra, gradually nar- rowed from base to apex, sparsely setose in an- terior %; mesosternum between coxae subequal in width to prosternal process. Pro femur with few moderately long setae; pro- tibia with spine field covering basal xlt^k of pos- terior face (Fig. 136); meso- and metatibia spi- DOYEN: SYSTEM ATICS OF El'SATIVS AND COS ISA III S 95 Figures 138-139. Posterior aspects of left forelegs. 138. Tibia of Eusattus difficilis. 139. E. productus. nose; tarsomere length ratios as follows: 5.4:1.7: 1.6:1.6:4.5 (pro-); 7.0:2.9:2.9:2.5:6.5 (meso-); 10.3:3.6:3.0:5.9 (meta). Measurements. — EL 10.3-14.1 mm; EW 8.7- 11.6 mm; PL 3.5-4.7 mm; PW 8.3-11.3 mm; BD 6.5-7.8 mm. Female. — Elytral declivity shining with tuber- cles interspersed with subcontiguous papillae Vi tubercle diameter; coxite length ratio to para- proct length = 0.36. Lectotype Female in MCZ. Type Localities.— Of both robust us and pos- tremis, San Clemente Island, California. Diagnosis. — Eusattus robustus is distin- guished from all other species by the broad epi- pleura which occupy the entire inflexed portion of the elytra. Distribution and Habitat. — Inhabits San Clemente Island, Los Angeles Co. and San Nic- olas Island, Ventura Co., California. On San Cle- mente Island, where the beetles are common, many habitats, excluding unstabilized sand dunes, are occupied. On San Nicolas Island, the beetles are largely restricted to the grassy central plateau and upper slopes and are much less abundant. Collections from San Clemente span the entire year; those from San Nicolas are from April and May, but adults are probably present throughout the year. Additional Material Examined. — California: Los Ange- les Co.: San Clemente Island: no additional data (51); 3A mi SW old airfield, VIII- 10- 1968 (3); China Point (7 corpses); Eagle Cyn., N of Gray, 1000-1500', IV-13-1980 (1); E slope Mt. Thirst, 111-22-1972 (6); vie. Mt. Thirst, Vlll-l 1-1968, IX- 10-1975, XI-1 1-1972 (5); Pyramid Head, IX- 1 1-1972 (7); West Cove, 111-21-1972, VI- 1 1-1971 (67); Wilson Cove, 01-23-1972 (4); 3 mi SE Wilson Cove, VI- 1 1-71 (5); 13.5 mi SE Wilson Cove, VI- 12- 1971 (5); Ventura County: San Nicolas Island: no additional data (8); NAS HQ area, V-5/6-1978 (27). Single individuals labeled Santa Barbara Island, San Miguel Island, Anacapa Island, and Santa Rosa Island are deposited in the LACM; they almost certainly represent labeling errors. Eusattus difficilis Species Group Head and protonum with duplex punctation, epistomum moderately to deeply and angulately emarginate medially (as in Fig. 59); slightly or not indented at lateral sutures; eyes reniform, constricted to about Vi greatest width by epistom- al canthus, dorsal and ventral lobes subequal; antenna subfiliform basally, gradually enlarged to 10th segment, apical segment asymmetrical with angulate apex; mentum subtrapezoidal, nearly flat; lateral margins nearly straight, then constricted before base; anterior angles acute, narrowly rounded; anterior border evenly, mod- erately deeply emarginate; submentum as wide as base of mentum; gular sutures strongly con- vergent, subcontiguous anteriorly. Pronotum strongly convex, moderately to broadly explanate laterally; lateral margins ar- cuate, anterior corners nearly right-angled, nar- rowly rounded; posterior corners acute, moder- ately produced posteriorly; hypomeron polished, glabrous except for dense submarginal fringe of long, projecting setae and few setae along ante- rior border; presternum before coxae much 96 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 141 shorter than prosternal process; prosternal pro- cess 1 .5 x wider behind than between coxae, flat, elongate, gradually tapering to rounded apex. Elytra 1.2-1.3 x wider than long, widest at about middle, very faintly to distinctly costulate; pseudepipleuron rounded, undefined; epipleuron gradually narrowed from base to apex, muri- cately punctate, setose in anterior %; mesoster- num between coxae narrower than prosternal process, deeply excavate anteriorly; metasternal suture about lh as long as metasternum. Femora bearing dorsal fringes of long, projecting setae; protibia very broadly triangular, outer margin slightly concave, with row of closely spaced or contiguous spinules reaching apex (Figs. 138, 139); apical process reaching approximately to apex of basal tarsomere. Metendosternite with arms fused with meta- notum apically, with mesocoxal inflections basally; spermatheca developed as 2 long unbranched, slender, convoluted tubules, ac- companied by 1-2 short tubules (Fig. 8) or as a pair of tubules, branched near the base (as in Fig. 9). Aedeagus bluntly rounded; spatulate pro- cesses of ovipositor straight, rounded apically; coxite length ratio to paraproct length = 0.34. Eusattus difficilis LeConte Eusattus difficilts LeConte, 1851:132; 1858:37; 1866:112 Lacordaire 1859:221; Horn 1870:294; 1883:305; 1894 423 (key); Casey 1908:70; Blaisdell 1921:213; 1943:193 Doyen 1977:5 (synonymy). Eusattus coquilletti Linell, 1899:180; Casey 1908:69. Eusattus convexus, Fall 1901:30, 166. Eusattus agnatus Casey, 1908:70. Eusattus compositus Casey, 1908:70. Eusattus coquilleti, Gebien 1938:284 (403); Papp 1961:116 (misspelling). Eusattus acutangulus, Doyen 1977:5. Eusattus congener, Doyen 1977:5. Broadly oval, convex black beetles with faintly costulate elytra. Male. — Frons set with large punctures 2-3 eye facets in diameter, separated by 1-2 puncture diameters, becoming denser, sometimes nearly confluent, and often coarser on epistomum and along epistomal suture; small punctures V3-V4 eye facet in diameter, separated by about 1 puncture diameter; epistomum faintly to moderately ru- gulose, entire or slightly indented at lateral su- tures. Antennal segment length ratios as fol- lows: 2.9:1.2:1.7:1.6:1.4:1.4:1.5:1.5:1.5:1.4:1.4: 1.4. Terminal segment 0.9-1.2 x broader than long; mentum about 1.7 x broader than long. Pronotum 2.2-2.3 x broader than long; disk with larger punctures slightly muricate, 2-3 eye facets in diameter, separated by 1-3 puncture diameters, becoming more strongly muricate, slightly coarser, and bearing very short, fine setae laterally; presternum and prosternal process punctato-rugulose, sparsely set with long, erect setae. Elytral disk weakly rugulose or eroded, depres- sions set with tubercles or very strongly muricate punctures about as large as or slightly larger than pronotal punctures; raised areas coalescing into 6-8 irregular, glabrous costulae, often very faint. Thoracic sternites and lateral quarters of abdom- inal sternites 1-3 sparsely set with long setae arising from muricate punctures, erect on thorax, declined on abdomen; medial areas of abdomi- nal sternites set with short, declined or appressed setae. Tarsomere length ratios as follows: 3.2:0.8: 0.8:0.8:2.4 (pro-); 4.6: 1.5:1.4:1 .0:2.8 (meso-); 5.8: 1.6:1.4:3.1 (meta-); tarsomeres subglabrous or with few very short, fine setae dorsally; terminal article with ventral lines of short, spiniform se- tae, becoming longer near apex. Female. — Differs as stated in generic and species group descriptions. Measurements. — EL 5.4-10.5 mm; EW 4.1- 8.4 mm; PL 1.8-3.4 mm; PW 3.9-7.9 mm; BD 2.8-6.0 mm. Holotype Female in MCZ. Type Localities.— Of difficilis, San Diego and Vallecitas, California; agnatus and coquilletti, Los Angeles Co., California; compositus, Oregon. Diagnosis.— Eusattus difficilis is extremely similar to E. convexus LeConte in body shape, size, and, superficially, in sculpturing. In convex- us, the spermatheca is of the type occurring in the reticulatus species group, with several short, thick tubules, while in difficilis it consists of 2- 3 very long, slender, tightly convoluted tubules, frequently accompanied by 2 shorter tubules. In addition, in difficilis the outer protibial margin has a row of subcontiguous spinules reaching the apex, while in convexus the spines are separated by 2-4 spine widths and extend no further than y» the distance to the apex. Variation. — Eusattus difficilis is geographi- cally variable in size and cuticular sculpturing. Individuals from cismontane California and Baja California and the western edge of the Mojave Desert have distinctly tuberculate elytra and more coarsely punctate pronota. East and north into Owens Valley, California, and southern Nevada. DOYEN: SYSTEMATICS OF EUSATTUS AND COMSA 11 1 \ 97 i ? :-'. m "»: Figure 140. Known distribution of Eusattus difficilis. elytral sculpturing becomes muricately punctate, pronotal punctation finer, and the setae on the dorsal metafemoral surface and on the abdom- inal sternites are longer. In Baja California south of the Sierra Martir, the elytra are shallowly and finely eroded, the erosions containing setigerous, somewhat muricate punctures. The thoracic ven- ter, abdominal sternites, and metafemoral setae are much longer than in cismontane California populations, and moderately long setae are pres- ent on the lateral edges of the pronotal disk. Body size averages largest in the Los Angeles basin and western Mojave Desert (6.2 mm < EL < 10.5 mm), is intermediate in the northeastern part of the range, and smallest in central Baja California (5.4 mm < EL < 7.3 mm). Distribution (Fig. 140) and Habitat. — E. difficilis occurs in diverse arid and semiarid hab- itats, from sea level to at least 2400 m elevation. It ranges from Inyo and Kern cos., California, and Nye Co., Nevada, in the north, through southern California to Bahia de Los Angeles, Baja California Norte, and El Desemboque in north- ern Sonora, Mexico. Most collections are from semiarid grassland or coniferous or deciduous woodland situations, often on sandy substrates, but clay or rocky soils may also support the bee- tles, as at McKittrick, Kern Co., California, and various localities in the Los Angeles basin. The collections from El Crucero and Bahia de Los Angeles, Baja California Norte, are from sand dunes. In this situation, the beetles may burrow into the sand at the bases of plants. On harder substrates, they shelter in litter beneath plants, in rodent burrows, or similar hiding places. The apparently isolated populations in north- ern Sonora (Puerto Penasco, El Desemboque) are phenetically very similar to those from Bahia de Los Angeles, and probably inhabit dune sands. Eusattus productus LeConte (Figure 141) Eusattus productus LeConte, 1858:20, 37; 1866:112; Horn 1870:295, 1883: 205; 1894:349, 423 (key); Casey 1908:67; Doyen 1977:6 (synonymy). Figure 141. Eusattus productus LeConte. 98 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 T~ " "■■'■■ i 1 - " ."■*" **T y ■' **.* *"^v**"".' T^*" /"*"-- / "*"'-->,.. / /' A """""••-.'' .'■' ,■• ) ■' r"""---. ; V <- ' / ^ s - \/^^ ♦ ■* P* S >. /, / \ \ ; - i \ • * \ :""•■■-. / O* • -\ ["-' "!'-■-- *o\ y "■ ' ' • % \ > % v U» / i-«0.39). 56. Ovipositor proportions (coxite length ratio to paraproct length): a. coxite moderate to long (>0. 28); b. coxite short (< 0.28). 57. Paramere shape: a. apically rounded; b. apically attenuate, pointed. 58. Spermathecal configuration: a. long, slender tubes; b. short, convoluted, irregular. 59. Spermathecal tube number: a. 4 (including state b. char. 58b); b. 2. 60. Spermathecal tube branching: a. unbranched or with irregular, apical branches; b. basally dichotomous. 61. Bursa copulatrix: a. absent; b. present. 102 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Appendix B Distribution of Character States3 Character 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 Conisattus rectus 1 1 1 1 1 1 1 2 2 1 1 1 1 2 2 1 1 2 1 Eusattus araneosus 1 2 1 1 2 1 1 2 1-2 arenarius 2 2 1 1 1 2 2 1 1 2 2 1 1 aridus 1 2 1 2 i 2 2 catalinensis 1 2 1 2 l 1 1 2 catavinus 1 2 1 2 1 1-2 [ 2 cedrosensis 1 2? 1 — — 1 1 2 ceralboensis 1 2 1 2 1 1 2 clllatoides 2 2 1 1 1 2 2 2 1 2 1 1 ciliatus 2 2 l 1 1 2 2 1 1. 2 1 1 cienegus 1 2 (1)2 1 2 1 2 1 1 2 convexus 1 2 1 2 1 1 I 1 costatus 1 2 1 2 i 2 2 crypticus 1 1 2 1 1 2 1 1 depressus 1 1 1 1 1 1 2 1 1 1 difficilis 1 1 (D2 1 1 1 2 1 2 1 1 dilatatus 2 1 2 2 1 2 1-2 2 2 1 2 1 1 dubius abditus 1-2 1 1 2 1 1 2 I 1 d. anzonensis 1 1 2 1 1 2 I 1 d. dubius 1 1 2 1 1 2 I 1 d. setosus 1-2 1 1 2 1 1 2 1 1 erosus erosus 1 2 2 1 1-2 2 e. manuelis 1 2 2 1 1 2 franciscanus 1 2 2 1 1 2 hirsutus 1 2 2 2 1 2 2 2 I 1 laevis 1 2 2 1 i ; I 2 mexicanus 1 2 1 2 2 l 2 i i minimus 1 2 2 i 2 1 mur. diabloensis 1 2 2 1 2 (1)2 2 1 2 1 mur. muncatus i 2 2 i 2 (1)2 2 1 2 1 nitidipennis 1 ^ 2 1 1 1 obliteratus 1 2 T ] 1 1 pallidus adustus 2 1 1 2 2 1 1 1-2 1 1 p. pallidus 2 1 1 2 2 1 1 1-2 1 1 phreatophilus 2 i 2 1 1 1 2 2 1 2 2 1 planulus 1 2 2 l 2 2 politus cruzensis 1 1 2 1 2 1 p. politus 1-2 1 2 1 1 1 2 5 1 pons 1 2 2 1 2 1 2 productus 1 1 2 1 1 2 1 2 1 puberulus 2 2 1 i ~\ 2 i 2 2 2 1 1 1 reticulatus 1 ■) 2 1 1 2 robustus 1 1 2 1 2 2 1 rudei 1 1 T 1 2 1 secutus 1 1 1 1 1 2 1 1 1 venosus 1 2 1 2 ~l 1 2 1 1 2 vizcainensis 2 1 1 1 1 2 2 1 ! 1 1-2 1 1 1 ' Primitive state indicated by 1. derived state by 2; 1-2 indicates that both states occur. Numbers in parentheses indicate less common states. Missing data indicated by bars. DOYEN: SYSTEMATICS OF EUSATTUS AND COX/SAULS 103 Appendix B Continued Character 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 Conisattus rectus 2 1 1 1 1 1 l l 1 1 1 1 1 1 1 1 1 1 2 Eusattus araneosus 1 2 1 2 1 1 1 1 1 arenarius 2 1 1 1 2 2 2 2 2 2 aridus 1 2 1 2 1 1 1 1-2 2 1 1 1 1 catalinensis 1 2 1 2 2 2 2 1 1 1 1 catavinus 1 2 2 2 1 1 1 1 cedrosensis 1 2 1 2 1 1 2 1 1 1 1 — ceralboensis 1 2 1 2 2 2 2 1 1 1 1 ciliatoides 2 1 1 1 2 2 2 2 2 2 ciliatus 2 1 1 1 2 2 2 2 2 2 cienegus 1 2 1 1 2 2 2 1 1 1 1 convexus 1 2 1 1 1-2 1 1 1 1 1(2) costatus 1 2 1 1-2 1 1 1 1-2 2 1 1 1-2 1 crypticus 2 1 1 1 2 1 2 1 2 2 depressus 1 1 1 1 1 1 1 1 difficilis 2 1 1 1 1-2 1 2 2 dilatatus 2 2 1 1 2 1 2 2 dubius abditus 2 1 1 1 1 1-2 2 1-2 1-2 d. arizonensis 2 1 1 1 1 1-2 2 1-2 1 d. dubius 2 1 1 1 1 1-2 2 1-2 1 d. setosus 2 1 1 1 1 1-2 2 1(2) 2 erosus erosus 1 2 2 2 1 1 1 e. manuelis 1 2 1-2 2 1 1 1 franciscanus 1 2 1 2 1 1 1 hirsutus 2 1 1 1 2 2 2 1 laevis 1 1 1 1 1 1 1 mexicanus 1 2 1 2 1 1 1 minimus 1 1 1 1 1-2 2 2 mur. diabloensis 2 1-2 1 1 1-2 2 2 2 2 mur. muricatus 2 1-2 1 1 (1)2 2 2 2 2 (1)2 nitidipennis 1 1 1 1 1 1 1 obliteratus 1 1 1 1 L 1 1 1 2 1 pallidus adustus 1-2 1 1 1 1 2 2 p. pallidus 2 1 1 1 1 2 2 2 phreatophilus 2 1 1 1 1 2 2 2 2 planulus 1-2 1-2 1 1-2 1 2 2 2 politus cruzensis 1 1 1 1 1 2 1-2 1 p. politus 1(2) 1 1 1 1 2 1-2 1 pons 1 2 2 1 2 2 2 2 2 1 1 1 product us 2 1 1 1 1-2 2 1-2 puberulus 1 1 1 1 2 1 1 1-2 2 2 2 2 reticulatus 1 2 1 I 1 2 2 2 1 1 1 robustus 2 1 1 1 2 1 1 1 1 rudei 1 1 1 1 2 2 2 secutus 1 1 1 1 1 1(2) 2 venosus 1 2 1 1 1 2 2 1-2 2 l 1 1 1 vizcainensis 2 1 1 1 1 1 1 1 1 1 1 2 2 2 2 104 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 141 Appendix B Continued Character 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 Comsattus rectus 1 2 1 1 2 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 Eusattus araneosus 1 1 1 2 1 1 arenarius 2 1 2 2 i 1 1 2 aridus 1 2 2 2 1 2 2 1 catalinensis 1 1 1 2 1 catavinus 1 2 2 2 1 cedrosensis 1 — — — 1 — 1 ceralboensis 1 1 1 2 1 ciliatoides 2 1 1 2 1 2 2 1 1 2 ciliatus 2 1 1 2 2 2 2 1 1 2 cienegus 1 2 2 1 2 2 1 convexus K2) 2 2 1 1(2) 2 1 costatus 1 (1)2 2 1 1(2) 2 1 crypticus 2 1(2) 2 2 2 1 1 depressus 1 2 2 2 2 1 2 2 1 2 2 i difficilis 2 1 2 1 1-2 i 1 2 2 i dilatatus 2 1 2 2 2 2 2 2 2 1 2 2 1 dubius abditus 1-2 1 1 1 1 1 2 d. ariionensis 1 1(2) 2 1 1 1 2 d. dubius 1 1(2) 2 1 1 1 2 d. setosus 2 1(2) 2 1 1 1 2 erosus erosus 1 2 2 1 1 2 2 e. manuelis 1 2 2 1 1 2 2 franciscanus 1 2 2 1 1 2 i hirsutus 1 1 1 2 1 1 — laevis 1 2 2 1 1 2 2 mexicanus 1 2 2 1 1 2 2 minimus 2 2 2 2 2 1 1 7 mur. diabloensis 2 2 2 2 1 2 2 mur. muricatus 2 2 1(2) 2 2 1 2 2 nitidipennis 1 1 1 1 1 1 2 obliieralus 1 1 2 1 1 1 2 pallidus adustus 2 2 2 2 2 2 1 2 1 p. pallidus 2 2 2 2 2 2 1 2 1 phreatophilus 2 2 2 2 1 1 2 2 2 planulus 2 2 1 1 1 1 2 politus cruzensis 1 1 2 1 1 1 1 p. politus 1 1(2) (1)2 1 2 1 1 1 1-2 1 pons 1 2 2 1 1 1 2 2 product us 1-2 1 1 1 1 2 1-2 1 2 2 1 puberulus 2 2 2 2 2 1 2 2 reticulatus 1 2 1 1 1 1 2 robustus 1 2 1 1 1 1 1 1 rudei 2 1(2) 2 2 1 1 1 1 1 2 secutus 2 2 2 2 i 1 1 1 2 1 2 2 1 venosus 1 2 2 1 1 1 2 1 2 2 vizcainensis 2 1 1 2 2 i 1 1 1 2 1 UH l^GS .